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Parasaccogaster rhamphidognatha (Cohen, 1987) Figs. 23, 24, 26, Tables 1, 2, 7 Saccogaster rhamphidognatha Cohen 1987: 1, fig. 1 (type locality: Gulf of Mexico). Saccogaster rhamphidognatha: Nielsen et al. 1999: 109. Material examined (1 specimen, 60 mm SL). Holotype: LACM 44189 - 1, male, 60 mm SL, Gulf of Mexico, Mississippi delta, 29 �� 13.5 ���N, 88 �� 16.5 ���W, R/V Alaminos, st. 69 A 13-43, 210 m, 4���16 Oct. 1969. Diagnosis. Parasaccogaster rhamphidognatha differs from the other two species of the genus by the antrorse, hook-like projection at the postero-ventral angle of the maxillary, a median spine on frontals above and behind eyes and the low number of pectoral fin rays (12). Also the following combination of characters is diagnostic: A strong, ethmoidal spine; pectoral peduncle prolonged and adnate to body; 2���3 slightly developed rakers on anterior gill arch; palatines with a single tooth row; gill filaments on anterior arch short, 1���2 times length of developed rakers; precaudal vertebrae 20 and total vertebrae 58; fin rays in dorsal 77, anal 49 and pectoral 12. Similarity. Parasaccogaster rhamphidognatha is most similar to P. n o r m a e with hidden opercular spine, nasal chamber not sooty black and distinctly more anal fin rays and vertebrae than found in P. melanomycter. P. rhamphidognatha differs from P. normae by the shorter gill filaments on anterior arch (1���2 times length of long rakers vs. 4���5 times), fewer pectoral fin rays (12 vs. 17���19), presence of ethmoidal spine (vs. spine absent) and a single median spine on frontal (vs. pair of small spines). Description. The principal meristic and morphometric characters are shown in Table 7. Head and body elongate and compressed with blunt snout and tapering tail. Scales absent. Lateral line obsolete. Dorsal fin origin above gill slit, anal fin origin well behind midpoint of fish and pectoral fins very short ending 1 / 5 to origin of anal fin. Anterior gill arch with 2���3 flat, spiny plates on upper branch, a short, slightly prolonged raker in angle and lower branch with 1���2 similar sized rakers followed by six spiny plates. Gill filaments short. Two pseudobranchial filaments. The intromittent organ is provided with a 6 mm long penis indicating a ripe male. Axial skeleton (from radiographs): Number of precaudal vertebrae 20. Anterior neural spine one third the length of second spine. Neural spines gradually decreasing in length. Neural spines 6���8 with slightly blunt tips. Tips of all other neural and haemal spines pointed. Parapophyses on vertebrae 4���20. Ribs obsolete. Dentition: Palatines with small, pointed teeth in a single row. Vomer boomerang-shaped with band of small, pointed teeth. Premaxillaries and vomer with an outer, narrow band of granular teeth and an inner row with larger, needle-like teeth. Head morphology: Head profile not concave above eyes. Head with single, strong, median, subdermal spine on frontal plate above and behind eyes, a strong median, subdermal, ethmoidal spine in front of eyes and a broad, blunt, sub-dermal spine above each eye. Anterior nostril placed close to upper lip, posterior nostril about three times as large, lunate-shaped, placed at half way distance to small eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thick, loose. Head with blunt snout, sunk-in eye covered by window of translucent skin. Upper jaw ends well behind eye with posterior margin strongly vertically expanded, somewhat hook-shaped. Flat opercular spine hidden below thick skin. No subdermal preopercular spine. Gill opening ending at about level of dorsal margin of pectoral peduncle. Otoliths: Not extracted from unique holotype. Coloration: Head and body pale. Vertical fins brownish distally. Biology and distribution (Fig. 26). Parasaccogaster rhamphidognatha is only known from the holotype caught at a depth of 210 m in the Gulf of Mexico., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 32-33, DOI: 10.5281/zenodo.208677, {"references":["Cohen, D. M. (1987) Notes on the bythitid fish genus Saccogaster with a new species from the Gulf of Mexico. Contribution in Science, 385, 1 - 4","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp."]}

Saccogaster parva Cohen & Nielsen, 1972 Figs 13, 14, 26, Table 1, 2, 5 Saccogaster parva Cohen & Nielsen 1972: 459, fig. 5 (type locality: off southern Brazil). Saccogaster parva: Nielsen et al. 1999: 110. Material examined (1 specimen, 58 mm SL). Holotype: ZMH 25268, female, 58 mm SL, off southern Brazil, 24 �� 21 ���S, 43 �� 54 ���W, R/V Walther Herwig, st. 90 / 68, bottom trawl, 500 m, 2 Mar. 1968. Diagnosis. Saccogaster parva differs from all other Saccogaster species in having 5���6 developed rakers on anterior gill arch and no spines on frontal plate and ethmoid. Also the following combination of characters is diagnostic: Skin thin and transparent; numerous neuromasts below translucent head skin arranged in five distinct clusters; scales absent; three developed rakers on anterior gill arch 3���4 times length of gill filaments; palatines with one row of teeth; antero-ventrally directed spine on lower angle of preoperculum; prolonged pectoral peduncle; precaudal vertebrae 16 and total vertebrae 54; fin rays in dorsal 91, anal 64 and pectoral 14. Similarity. Saccogaster parva and S. nikoliviae are the only species with numerous, developed neuromasts on the head, but they differ in other characters such as number of developed gill rakers 6 vs. 3 in S. nikoliviae, absence of scales vs. present in S. nikoliviae and no spines behind eyes or on ethmoid vs. present in S. nikoliviae. Description. The principal meristic and morphometric characters are shown in Table 5. Body compressed and relatively short. Scales absent. Lateral line continuous, originating above preopercle descending gradually to midline over anus and ending close to basis of caudal fin; ca. 20 lateral line papillae. Dorsal fin origin above proximal part of pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fins end halfway to anus. Pectoral peduncle prolonged. Head compressed with blunt snout. Opercular spine flat, preopercular spine hidden. Posteror, vertically expanded part of maxillary strongly sheathed dorsally. Anterior gill arch (Fig. 14 E) with two knobs on upper branch, a developed raker in angle, and lower branch with 5���6 developed rakers followed by 6���8 knobs. Pseudobranchial filaments 2. Ovaries with developing eggs. Axial skeleton (from radiographs): Number of precaudal vertebrae 16. Anterior neural spine 1 / 3 length of second spine. Neural spines 2���5 decreasing in length and with pointed tips. Neural spines 6���10 with blunt tips and all remaining neural and haemal spines with pointed tips. Parapophyses developed on vertebrae 7���16. Pleural ribs on vertebrae 2���11 and epipleural ribs seen on vertebrae 3���6. Dentition: Vomer boomerang-shaped with about six fangs and a scattering of smaller teeth. Palatines with one row of fang-like teeth. Dentaries with an irregular row of about 15 fangs and an outer narrow band of shorter teeth. Premaxillaries with a narrow band of small, granular teeth and five needle-like teeth at the symphysis Head morphology (Fig. 14 A���D): Head profile straight above eyes. Head without spines on frontal plate and on ethmoid, but blunt, sub-dermal spine above eyes. Fig. 14 B shows a conspicuous thickening of the frontal at the location where spines would be expected, indicating that larger specimens might in fact have small frontal spines. Also a broad hump is indicated on the snout at the position where a (cartilaginous) spine might be expected. Anterior nostril placed close to upper lip; posterior nostril moderately large with anterior flap, placed close to eye. Head pores: 2 supraorbital pores near tip of snout, 1 anterior infraorbital pore below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, tight. Total of 17���18 diamond-shaped, light-colored neuromasts visible below transparent skin arranged in five clusters: 4 infraorbital along upper lip and curving around eye, 2 pairs between eyes, one of them slightly in front of eyes, 6 supraorbital along line above opercle and behind eyes, 2���3 mandibular along jaw, 3 on the lower preopercle. Head with dusky background pigmentation chiefly around neuromasts. Upper jaw ends well behind eye; posterior end of maxilla vertically expanded. Opercular spine pointed but flat, subdermal, not reaching hind margin of opercle. Small, subdermal anterior-ventrally pointed spine at lower angle of preoperculum. Otolith: Dissolved. Coloration: Violet when caught. Biology and distribution (Fig. 26). Caught on the bottom on the upper continental slope at a depth of 500 m. Known only from the holotype off southern Brazil., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 19-20, DOI: 10.5281/zenodo.208677, {"references":["Cohen, D. M. & Nielsen, J. G. (1972) A review of the viviparous ophidioid fishes of the genus Saccogaster. Proceedings of the Biological Society of Washington, 85 (39), 445 - 468.","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp."]}

Key to the species of Saccogaster 1 Scales present on body................................................................................ 2 - Scales absent from body................................................................................ 6 2 Opercular spine free, with 3 prongs; pectoral fin rays 22; dorsal fin rays 92.................................. hawaii - Opercular spine free or hidden, with single prong; pectoral fin rays 16–19; dorsal fin rays 80–88....................... 3 3 Gill filaments on anterior arch short (1 / 3 – 1 / 5 times length of developed rakers)................................... 4 - Gill filaments on anterior arch about equal to or longer than developed gill rakers.................................. 5 4 Scales present on predorsal; 12–13 precaudal vertebrae; no brown stripe on head; numerous neuromasts...... nikoliviae n.sp. - Scales absent on predorsal; 21 precaudal vertebrae; broken brown stripe on sides of head; no neuromasts....... brayae n.sp. 5 Gill filaments on anterior arch distinctly longer than developed gill rakers; opercular spine free; palatines with several tooth rows; dorsal fin rays 83–85; posterior infraorbital pores 2, posterior supraorbital pore 1, posterior mandibular pore 1....... maculata - Gill filaments on anterior arch 0.5–1 times length of developed gill rakers; opercular spine hidden; palatines with a single tooth row; dorsal fin rays 86–88; no posterior infraorbital pores, no posterior supraorbital pore, no posterior mandibular pore..................................................................................................... staigeri 6 Developed gill rakers on anterior arch 5–6; pectoral fin rays 14; numerous neuromasts........................... parva - Developed gill rakers on anterior arch 3; pectoral fin rays 19–22; 0–3 neuromasts................................... 7 7 Skin thin; opercular spine free; spines on frontal plate visible, bony ridges above eyes; distance from base of pelvic fins to ori- gin of anal fin 46.0% SL; vomer and premaxillaries with fangs; no posterior mandibular pore.................... horrida - Skin thick; opercular spine hidden; spines on frontal plate sub-dermal, no bony ridge above eyes; distance from base of pelvic fins to origin of anal fin 32.0–39.0 % SL; no fangs, dentition granular; large posterior mandibular pore......... tuberculata

Saccogaster horrida n. sp. Figs. 7, 8, 25, Tables 1, 2, 4 Hastatobythites arafurensis (non Machida): Hoese & Gates 2006: 573. Material examined (1 specimen, 59 mm SL). Holotype: NMV A 29718 -019, female, 59 mm SL, off northwestern Australia, Leveque L 27 transect, 14 ° 51 ’ 12 ’’S, 121 ° 25 ’ 53 ’’E to 14 ° 50 ’ 43 ’’S, 121 ° 27 ’01’’E, R/V Southern Surveyor, beam trawl, 403 – 396 m, 2 July 2007. Diagnosis. Saccogaster horrida differs from all other Saccogaster species by the following combination of characters: Distinct pair of free spines on frontal plate above and behind eyes, bony ridge above eye and cartilaginous ethmoidal spine in front of eyes; no scales, thin, transparent skin, no neuromasts; long gill rakers on anterior arch 2–3 times length of gill filaments; palatines with 2–3 tooth rows; pectoral peduncle twice as long as high; precaudal vertebrae 14 and total vertebrae 56; fin rays in dorsal 84, anal 57 and pectoral 21; otolith length to height = 1.75, otolith length to colliculum length = 2.1, sulcus with long pointed anterior projection. Similarity. Saccogaster horrida seems most close to S. tuberculata, both lacking scales on the body. They differ from each other by S. horrida having a distinct frontal plate with a pair of well-developed spines (vs. no distinct plate and small sub-dermal spines), bony ridge above eye (vs. no bony ridge), vomer and premaxillaries with fangs (vs. no fangs), distance from base of pelvic fins to origin of anal fin 46 % SL (vs. 32–39 % SL) and ratio otolith length to height = 1.75 (vs. 2.1–2.2). Description. The principal meristic and morphometric characters shown in Table 4. Head and body elongate and compressed with tapering caudal part and blunt snout. Scales absent. Lateral line not visible. Dorsal fin origin above middle pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fin ending more than halfway to anus and with prolonged peduncle. Anterior gill arch (Fig. 8 D) with six spiny knobs on upper branch, a developed raker in angle and lower branch with two developed rakers with a knob in between followed by 11 knobs. Developed rakers on anterior arch 2–3 times length of gill filaments. Pseudobranchial filaments not found. Ovaries well developed. Axial skeleton (from radiographs): Number of precaudal vertebrae 14. Anterior neural spine one third length of second spine. Neural spines 2–4 long and depressed with pointed tips. Neural spines 5–14 shorter, with large basal parts and with pointed tips except for blunt in spines 6–9. Parapophyses developed on vertebrae 7–14. Pleural ribs on vertebrae 3–6 and epipleural ribs not seen. Dentition: Palatines with 2–3 irregular rows of small, pointed teeth. Vomer boomerang-shaped and with 2–3 irregular tooth rows; inner row with ten large, pointed teeth and outer rows with small, pointed teeth. Premaxillaries with 3–5 irregular rows of granular teeth except for 2–3 fangs at symphysis. Dentaries with 3–4 irregular rows of small, pointed teeth except for larger teeth in inner row. Head morphology (Fig. 8 A–C): Head profile concave above eyes. Head with pair of free spines on frontal plate above and behind eyes, bony ridge above eye, frontal plate with rough-textured surface below skin, median, sub-dermal cartilaginous ethmoidal spine in front of eyes and median bony ridge (3 mm long) on dorsum behind frontal plate. Anterior nostril placed close to upper lip and small posterior nostril placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 1 anterior infraorbital pore below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, tight. No neuromasts visible. Head with pigmentation patches around mouth and above eyes and opercular spine. Upper jaw ends well behind eye with posterior margin vertically expanded. Opercular spine pointed, almost reaching hind margin of operculum. No spine at lower angle of preoperculum. Otolith (Fig. 8 E–F): Otolith oval, moderately thick; otolith length to height = 1.75, otolith height to thickness = 1.9. Dorsal and ventral rims gently and regularly curved, smooth. Anterior and posterior tips regularly rounded resulting in a symmetrical outline of the otolith. Inner face slightly convex with centrally positioned, undivided oval, shallow sulcus with long and pointed anterior projection; otolith length to colliculum length = 2.1. Outer face convex. Coloration: After five years of preservation body uniformly brownish. Head with scattered, small, black pigment mainly dorsally and on jaws. Frontal plate with a few larger, black pigment spots. Eyes blue with whitish lens. Biology and distribution (Fig. 25). A viviparous species caught on the upper continental slope at a depth of 396–403 m off northwestern Australia. Known from a female with ovaries containing many eggs but no embryos. Etymology. The name “ horrida ” (meaning horrifying) refers to the spines and bony ridges above the eyes.

Parasaccogaster n.gen. Type species: Saccogaster normae Cohen & Nielsen, 1972 Diagnosis. A genus of the subfamily Bythitinae (Cohen & Nielsen 1978) characterized by the following combination of characters: Scales absent on head and body, head skin thick and loose; eyes sunk-in below transparent skin-window; gill opening at level of or below dorsal margin of pectoral peduncle; head width 7.7 ���12.0 % SL, maxilla vertically expanded posteriorly; a median or a pair of small bony spines on frontals covered by skin above and behind eyes; palatine teeth present; developed gill rakers on anterior arch 2���3, pseudobranchial filaments 2; males with stalked intromittent organ; vertical fins joined, anal fin origin behind midpoint of fish, pectoral fin radials slightly longer than high, peduncle adnate to body; pectoral fin rays 12���19; precaudal vertebrae 14���20. Similarity. The most similar genera are Hastatobythites and Saccogaster. Parasaccogaster differs from both genera by having thick skin with sunk-in eyes, pectoral peduncle adnate to body and gill opening ending at level with or below dorsal margin of pectoral peduncle. Etymology. Parasaccogaster refers to the similarity to Saccogaster., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on page 26, DOI: 10.5281/zenodo.208677, {"references":["Cohen, D. M. & Nielsen, J. G. (1972) A review of the viviparous ophidioid fishes of the genus Saccogaster. Proceedings of the Biological Society of Washington, 85 (39), 445 - 468.","Cohen, D. M. & Nielsen, J. G. (1978) Guide to the identification of genera of the fish order Ophidiiformes with a tentative classification of the order. NOAA Technical Report NMFS Circular, 417, 1 - 72."]}

Key to species of Parasaccogaster 1 Nasal chamber black; vertebrae 45; anal fin rays 37; head width 12.0 % SL............................. melanomycter - Nasal chamber not black; vertebrae 54–58; anal fin rays 49–54; head width 7.7–9.8 % SL............................ 2 2 Gill filaments on first gill arch short (1–2 times developed rakers); precaudal vertebrae 20; pectoral fin rays 12; strong ethmoi- dal spine................................................................................ rhamphidognatha - Gill filaments on first gill arch long (4–5 times developed rakers); precaudal vertebrae 14–16; pectoral fin rays 17–19; no eth- moidal spine................................................................................... normae

Saccogaster hawaii Cohen & Nielsen, 1972 Figs. 5, 6, 25, Tables 1, 2, 4 Saccogaster hawaii Cohen & Nielsen 1972: 455, figs. 1, 3 (type locality: off Maui, Hawaian Islands); Nielsen et al. 1999: 109. Material examined (1 specimen, 69 mm SL). Holotype: USNM 207355, female, 69 mm SL, off Maui, Hawaii, 21 ��00���N, 156 �� 47 ���W, R/V Townsend Cromwell, st. 40���62, bottom trawl, bottom temp. 15.9 ��C, 234 m, 19 Nov. 1968. Diagnosis. Saccogaster hawaii differs from all other Saccogaster species by the opercular spine ending in three prongs. Also the following combination of characters is diagnostic: A pair of sub-dermal spines on frontal behind and above eyes, median sub-dermal ethmoidal spine in front of eyes with forward pointed tip; three developed rakers on anterior gill arch hardly longer than the knobs, extremely small filaments on anterior gill arch; scales absent on predorsal and present on median part of body; thin, transparent skin; palatines with single row of teeth; elongate pectoral peduncle longer than high; precaudal vertebrae 18 and total vertebrae 54; fin rays in dorsal 92, anal 55 and pectoral 22. Similarity. Saccogaster hawaii seems most similar to S. brayae (see comparison above). Description. The principal meristic and morphometric characters are shown in Table 4. Head and body elongate and compressed with tapering caudal part and blunt snout. Scales absent from predorsal part and from band along bases of dorsal and anal fins. Anteriormost scales widely scattered, those more posteriorly positioned closely spaced and on rear of body regularly imbricate. Lateral line indistinct. Dorsal fin origin above distal part of pectoral peduncle, anal fin origin well behind midpoint of fish and pelvic fins inserted below hind margin of opercle, ending below middle of pectoral fin. Anterior gill arch (Fig. 6 D) with three barely developed rakers scarcely longer than the three spiny knobs on upper branch and the knobs on lower branch. Very short gill filaments on anterior arch. No pseudobranchial filaments observed. No intromittent organ developed so it is probably an immature female. A pair of elongate gonads visible through transparent skin. Axial skeleton (from radiographs): Number of precaudal vertebrae 18. Anterior neural spine very short. Neural spines 2���4 long, depressed and with pointed tips. Neural spines 5���18 with short, blunt spines. Parapophyses developed on vertebrae 7���18. Pleural ribs distinct on vertebrae 3���6. Dentition: Palatines with 10���15 sharp, pointed teeth in a single row. Vomer with 6 teeth similar to the longer premaxillary teeth plus a scattering of smaller teeth. Premaxillaries with band of small, granular teeth and near symphysis as series of 8 elongate, needlelike teeth in a widely spaced row. Dentaries with an irregular row of about 15 longer, pointed teeth and an outer narrow band of shorter teeth. Otolith: Not extracted from unique holotype. Head morphology (Fig. 6 A���C): A pair of forward directed sub-dermal frontal spines behind and above eyes, a large, median, sub-dermal ethmoidal spine in front of eyes with forward pointed tip and a blunt sub-dermal spine above each eye. Five faint neuromasts below eye curving upwards and two on snout inward of posterior nostril. Upper jaw ends well posterior to eye with posterior margin vertically expanded and not sheathed dorsally. Opercle with a flattened, antler-like, 3 ���pronged spine (Fig. 6 A). Coloration: Body pale with fine, brown pigmentation along the bases of dorsal and anal fins. Similar pigmentation on head, particularly on snout and between eyes, and an intense dark band from behind eye becoming more diffuse posteriorad. Biology and distribution (Fig. 25). Saccogaster hawaii is only known from the holotype caught at the upper continental shelf at a depth of 234 m off Maui, Hawaiian Islands., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 10-12, DOI: 10.5281/zenodo.208677, {"references":["Cohen, D. M. & Nielsen, J. G. (1972) A review of the viviparous ophidioid fishes of the genus Saccogaster. Proceedings of the Biological Society of Washington, 85 (39), 445 - 468.","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp."]}

Saccogaster brayae n. sp. Figs. 3, 4, 25, Table 1, 2, 4 Material examined (1 specimen, 36 mm, SL). Holotype: NMV A 29655 -002, male, 36 mm SL, off northwestern Australia, Barrow L 1 transect, 21 °00’ 49 ’’S, 114 ° 39 ’09’’E to 21 °00’ 47 ’’S, 114 ° 38 ’ 54 ’’E, R/V Southern Surveyor, Sherman sledge, 250–258 m, 10 June 2007. Diagnosis. Saccogaster brayae differs from all other Saccogaster species in having a broken brown stripe between eye and hind margin of opercle (except for S. hawaii), precaudal vertebrae 21 and total vertebrae 53, and becoming ripe at a very small SL (36 mm). Also the following combination of characters is diagnostic: A pair of free spines on frontal above and behind eyes, a large median sub-dermal ethmoidal spine in front of eyes with forward pointed tip; antero-ventrally directed spine at lower angle of preopercle; thin transparent skin; scales present on sides of body; three long gill rakers on anterior arch 4–5 times the length of gill filaments; palatines with 1–2 rows of teeth; pectoral peduncle longer than high; fin rays in dorsal 85, anal 52 and pectoral 18; otolith length to height = 1.9, otolith length to colliculum length = 3.1, sulcus without anterior projection. Similarity. Saccogaster brayae seems most similar to S. hawaii by the brown stripe behind eye, the forward pointed tip of the ethmoidal spine and the scaled body. They differ by S. brayae having opercular spine ending in one prong (vs. three prongs), by long rakers on anterior arch 4–5 times length of filaments (vs. long rakers and filaments almost equal in length), precaudal vertebrae 21 (vs. 18), pectoral fin rays 18 (vs. 22) and dorsal fin rays 85 (vs. 92). Description. The principal meristic and morphometric characters are shown in Table 4. Head and body elongate and compressed with tapering caudal part and blunt snout. Most scales rubbed off but scale-pockets and a few remaining scales (0.8 mm long) indicate squamation on sides of body. Lateral line not visible. Dorsal fin origin above proximal part of pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fin ending less than halfway to anus with peduncle a little longer than broad and free of body. Anterior gill arch with three spiny knobs on upper branch, a developed raker in angle and lower branch with two developed rakers with a knob in between followed by eight knobs. Developed rakers 4–5 times length of gill filaments (Fig. 4 D). Two small pseudobranchial filaments. The well developed testes and intromittent organ (Fig. 4 G) show that the specimen is a ripe male. Axial skeleton (from radiographs): Number of precaudal vertebrae 21. Anterior neural spine very short. Neural spines 2–4 long, depressed and with pointed tips. Neural spines 5–20 short and with large basal parts, length of spines increasing posteriorad. Parapophyses developed on vertebrae 6–20. Pleural ribs distinct at vertebrae 1–8, but pleural ribs on remaining precaudal vertebrae and presence of epipleural ribs are questionable. Dentition: Palatines with about 25 pointed teeth in 1–2 rows; larger teeth in inner row. Vomer boomerangshaped with about 25 pointed teeth in 2–3 irregular rows; six longer teeth in inner row. Premaxillaries with granular teeth except for five fangs at symphysis. Dentaries with 3–4 irregular tooth rows, outer rows with granular teeth and inner row with fangs. Head morphology (Fig. 4 A–C): Head profile concave above eyes. Head with pair of free spines as an integrated part of a transverse rim on frontal above and slightly behind eyes and a large, median sub-dermal ethmoidal spine in front of eyes with forward pointed tip. Anterior nostril placed close to upper lip and small posterior nostril placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, tight. No neuromasts visible. Head with dusky background pigmentation, broken brown stripe between eye and hind margin of opercle and broken brown pigmentation patches below and behind eye. Upper jaw ends well behind eye with posterior margin vertically expanded. Opercular spine pointed, just reaching hind margin of operculum. Antero-ventrally directed spine at lower angle of preoperculum. Otolith (Fig. 4 E–F): Otolith oval, moderately thick; otolith length to height = 1.9, otolith height to thickness = 1.9. Dorsal and ventral rims gently curved, dorsal rim broadly undulating. Anterior and posterior tips regularly rounded resulting in a regular oval appearance of the otolith. Inner face flat with centrally positioned, undivided, small, oval, shallow sulcus without anterior projection. Otolith length to colliculum length = 3.1. Outer face convex. Coloration: After five years of preservation body uniformly brownish with faint dark pigmentation dorsally. Head with dusky, dense pigmentation and a dark brown stripe from eye to hind margin of opercle. Larger, dark pigment spots on frontal plate seen through skin on dorsum. Eyes blue with yellow lens. Biology and distribution (Fig. 25). A viviparous species occurring on the upper continental slope. Known from a 36 mm male with well developed testes and intromittent organ, caught off northwestern Australia organ (Fig. 4 G), caught at a depth of 250– 258 m. Etymology. Named after Dianne Bray, MNV, for her support with material for this revision.

Saccogaster horrida n. sp. Figs. 7, 8, 25, Tables 1, 2, 4 Hastatobythites arafurensis (non Machida): Hoese & Gates 2006: 573. Material examined (1 specimen, 59 mm SL). Holotype: NMV A 29718 -019, female, 59 mm SL, off northwestern Australia, Leveque L 27 transect, 14 �� 51 ��� 12 ������S, 121 �� 25 ��� 53 ������E to 14 �� 50 ��� 43 ������S, 121 �� 27 ���01������E, R/V Southern Surveyor, beam trawl, 403 ��� 396 m, 2 July 2007. Diagnosis. Saccogaster horrida differs from all other Saccogaster species by the following combination of characters: Distinct pair of free spines on frontal plate above and behind eyes, bony ridge above eye and cartilaginous ethmoidal spine in front of eyes; no scales, thin, transparent skin, no neuromasts; long gill rakers on anterior arch 2���3 times length of gill filaments; palatines with 2���3 tooth rows; pectoral peduncle twice as long as high; precaudal vertebrae 14 and total vertebrae 56; fin rays in dorsal 84, anal 57 and pectoral 21; otolith length to height = 1.75, otolith length to colliculum length = 2.1, sulcus with long pointed anterior projection. Similarity. Saccogaster horrida seems most close to S. tuberculata, both lacking scales on the body. They differ from each other by S. horrida having a distinct frontal plate with a pair of well-developed spines (vs. no distinct plate and small sub-dermal spines), bony ridge above eye (vs. no bony ridge), vomer and premaxillaries with fangs (vs. no fangs), distance from base of pelvic fins to origin of anal fin 46 % SL (vs. 32���39 % SL) and ratio otolith length to height = 1.75 (vs. 2.1���2.2). Description. The principal meristic and morphometric characters shown in Table 4. Head and body elongate and compressed with tapering caudal part and blunt snout. Scales absent. Lateral line not visible. Dorsal fin origin above middle pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fin ending more than halfway to anus and with prolonged peduncle. Anterior gill arch (Fig. 8 D) with six spiny knobs on upper branch, a developed raker in angle and lower branch with two developed rakers with a knob in between followed by 11 knobs. Developed rakers on anterior arch 2���3 times length of gill filaments. Pseudobranchial filaments not found. Ovaries well developed. Axial skeleton (from radiographs): Number of precaudal vertebrae 14. Anterior neural spine one third length of second spine. Neural spines 2���4 long and depressed with pointed tips. Neural spines 5���14 shorter, with large basal parts and with pointed tips except for blunt in spines 6���9. Parapophyses developed on vertebrae 7���14. Pleural ribs on vertebrae 3���6 and epipleural ribs not seen. Dentition: Palatines with 2���3 irregular rows of small, pointed teeth. Vomer boomerang-shaped and with 2���3 irregular tooth rows; inner row with ten large, pointed teeth and outer rows with small, pointed teeth. Premaxillaries with 3���5 irregular rows of granular teeth except for 2���3 fangs at symphysis. Dentaries with 3���4 irregular rows of small, pointed teeth except for larger teeth in inner row. Head morphology (Fig. 8 A���C): Head profile concave above eyes. Head with pair of free spines on frontal plate above and behind eyes, bony ridge above eye, frontal plate with rough-textured surface below skin, median, sub-dermal cartilaginous ethmoidal spine in front of eyes and median bony ridge (3 mm long) on dorsum behind frontal plate. Anterior nostril placed close to upper lip and small posterior nostril placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 1 anterior infraorbital pore below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, tight. No neuromasts visible. Head with pigmentation patches around mouth and above eyes and opercular spine. Upper jaw ends well behind eye with posterior margin vertically expanded. Opercular spine pointed, almost reaching hind margin of operculum. No spine at lower angle of preoperculum. Otolith (Fig. 8 E���F): Otolith oval, moderately thick; otolith length to height = 1.75, otolith height to thickness = 1.9. Dorsal and ventral rims gently and regularly curved, smooth. Anterior and posterior tips regularly rounded resulting in a symmetrical outline of the otolith. Inner face slightly convex with centrally positioned, undivided oval, shallow sulcus with long and pointed anterior projection; otolith length to colliculum length = 2.1. Outer face convex. Coloration: After five years of preservation body uniformly brownish. Head with scattered, small, black pigment mainly dorsally and on jaws. Frontal plate with a few larger, black pigment spots. Eyes blue with whitish lens. Biology and distribution (Fig. 25). A viviparous species caught on the upper continental slope at a depth of 396���403 m off northwestern Australia. Known from a female with ovaries containing many eggs but no embryos. Etymology. The name ��� horrida ��� (meaning horrifying) refers to the spines and bony ridges above the eyes., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 12-14, DOI: 10.5281/zenodo.208677, {"references":["Hoese, D. F. & Gates, J. E. (2006) Bythitidae In, Beesley, P. L. & Wells, A. (Eds.) Zoological Catalogue of Australia. Volume 35. ABRS & CSIRO Publishing, Australia, pp. 567 - 574."]}

Saccogaster nikoliviae n. sp. Figs.11, 12, 25, Tables 1, 2, 5 Material examined (3 specimens, 50���69 mm SL). Holotype: NMV A 29674 -010, female, 69 mm SL, off northwestern Australia, Imperieuse L 23 transect, 17 �� 31 ��� 44 ������S, 118 �� 50 ��� 37 ������E to 17 �� 32 ��� 30 ������S, 118 �� 50 ��� 31 ������E, R/V Southern Surveyor, beam trawl, 407 ��� 403 m, 15 June 2007. Paratypes: CSIRO H 6602 -01, female, 57 mm SL (same data as for holotype). NMV A 29670 -021, male, 50 mm SL, off northwestern Australia, Hedland L 22 transect, 18 �� 34 ��� 12 ������S, 117 �� 27 ��� 52 ������E to 18 �� 34 ���04������S, 117 �� 28 ��� 38 ������E, R/V Southern Surveyor, beam trawl, 405 ��� 401 m, 14 June 2007. Diagnosis. Saccogaster nikoliviae differs from all other Saccogaster species by presence of predorsal scales and by the following combination of characters: A more or less distinct pair of frontal spines above and behind eyes and a median, sub-dermal, cartilaginous, ethmoidal spine in front of eyes; numerous neuromasts below translucent head-skin arranged in four distinct clusters; scales on body; three long gill rakers on anterior arch 3���5 times length of filaments; pectoral peduncle prolonged; palatines with 2���3 tooth rows; an antero-ventrally directed spine on lower angle of preoperculum; precaudal vertebrae 12���13 and total vertebrae 52���53; fin rays in pectoral 17���18, dorsal 80���84 and anal 48���52; otolith length to height = 1.9, otolith length to colliculum length = 2.8���3.1. Similarity. Saccogaster nikoliviae and S. parva are the only species with numerous developed neuromasts on the head. They differ in number of developed gill rakers (3 vs. 6 in S. parva), presence of scales vs. none in S. parva, and a pair of spines behind eyes vs. none in S. parva. Description. The principal meristic and morphometric characters are shown in Table 5. The description is based on the holotype; differences from the two paratypes are mentioned in brackets. Body moderately compressed and elongate with tapering caudal part and blunt snout. Body covered with thin, overlapping scales on sides and on predorsal; length of scale from above pectoral fin 0.8 mm. No scales on head and abdomen. Lateral line indistinct (observed close to dorsal margin of body ending above tip of pectorals). Dorsal fin origin above proximal part of pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fins ending half way to anus with peduncle not attached to body. Pelvic fins ending below anterior part of pectoral fins. Anterior gill arch (Fig. 12 F) with three (4) knobs on upper branch, a long raker in angle and lower branch with two long rakers with a knob in between followed by seven (10) knobs; all knobs and long rakers with small densely placed spines. Long rakers on anterior gill arch 3 (5) times length of filaments. Ca. 35 very small gill filaments. Two thin pseudobranchial filaments. Axial skeleton (from radiographs): Number of precaudal vertebrae 12���13. Anterior neural spine one fourth the length of second spine. Neural spines 4 to 8 ��� 9 gradually decreasing in length. Pointed tips on vertebrae 2 to 4 ��� 5, blunt tips on vertebrae 6 to 10 ��� 11 and the remaining vertebrae with neural and haemal spines pointed. Vertebrae 3���7 with depressed tips and vertebrae 3 to 7 ��� 8 with enlarged basal parts. Parapophyses developed on vertebrae 7���13, pleural ribs on vertebrae 3 to 7 ��� 8 and epipleural ribs on vertebrae 3���10. Dentition: Vomer boomerang-shaped with 5���10 fangs in inner row and 20���25 small, close-set teeth in 2���3 outer rows. Palatines with about 10 fangs in inner row and many small teeth in 1���2 outer rows. Dentaries with 10���20 fangs in inner row and many, small teeth in 2���3 outer rows. Premaxillaries with 2���3 long fangs near symphysis and with granular teeth in several, irregular rows further back. Head morphology (Fig. 12 A���D): Head profile concave above eyes. Head with pair of small free, closely set spines on frontal above and behind eyes, sub-dermal surface of frontal plate very uneven, a median, subdermal, cartilaginous, ethmoidal spine in front of eyes and a blunt sub-dermal spine above eyes. Anterior nostril with a low rim placed close to upper lip; posterior nostril a mere hole placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, tight. Total of 17���18 diamond-shaped, light-colored neuromasts visible below transparent skin arranged in four cluster: 5���6 infraorbital along upper lip and curving around eye, 2 pairs between eyes, 6 supraorbital along line above opercle and behind eyes, 4 mandibular along jaw. Upper jaw ends well behind eye; posterior end of maxilla vertically expanded. Opercular spine pointed but flat, free, not reaching hind margin of opercle. Small, subdermal anterior-ventrally pointed spine at lower angle of preoperculum. Otolith (Fig. 12 E, G���H): Otolith oval, moderately thick; otolith length to height = 1.9, otolith height to thickness = 2.2. Dorsal and ventral rims gently and regularly curved, smooth. Anterior and posterior tips regularly rounded resulting in a symmetrical outline of the otolith. Inner face slightly convex with centrally positioned, undivided oval, shallow sulcus without anterior projection; otolith length to colliculum length = 2.8���3.1. Outer face convex. Coloration: Head and body light brown. Head with dusky background coloration. Eyes black with lighter lens. Biology and distribution (Fig. 25). A viviparous species occurring on the upper continental slope. The holotype and a 57 mm SL paratype are females with extended ovaries and eggs up to 0.5 mm in diameter; no embryos were observed. The 50 mm SL paratype is a male with ripe testes and well developed intromittent organ. Known from three specimens caught off northwestern Australia in bottom fishing gear at depths between 401 and 407 m. Etymology. The new species is named after two of the senior author���s grandchildren: Nikolaj and Olivia., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 16-18, DOI: 10.5281/zenodo.208677

Parasaccogaster n.gen. Type species: Saccogaster normae Cohen & Nielsen, 1972 Diagnosis. A genus of the subfamily Bythitinae (Cohen & Nielsen 1978) characterized by the following combination of characters: Scales absent on head and body, head skin thick and loose; eyes sunk-in below transparent skin-window; gill opening at level of or below dorsal margin of pectoral peduncle; head width 7.7 –12.0 % SL, maxilla vertically expanded posteriorly; a median or a pair of small bony spines on frontals covered by skin above and behind eyes; palatine teeth present; developed gill rakers on anterior arch 2–3, pseudobranchial filaments 2; males with stalked intromittent organ; vertical fins joined, anal fin origin behind midpoint of fish, pectoral fin radials slightly longer than high, peduncle adnate to body; pectoral fin rays 12–19; precaudal vertebrae 14–20. Similarity. The most similar genera are Hastatobythites and Saccogaster. Parasaccogaster differs from both genera by having thick skin with sunk-in eyes, pectoral peduncle adnate to body and gill opening ending at level with or below dorsal margin of pectoral peduncle. Etymology. Parasaccogaster refers to the similarity to Saccogaster.

Saccogaster tuberculata (Chan, 1966) Figs. 17, 18, 25, Table 1, 2, 6 Barbuliceps tuberculatus Chan 1966: 4, figs. 1���2 (type locality: South China Sea). Saccogaster tuberculata: Cohen 1987: 3; Nielsen et al. 1999: 110. Material examined (18 specimens, 84���160 mm SL). Holotype: BMNH 1965.11. 6.1, male, 106 mm SL, South China Sea, 6 ��01��� 48 ������N, 109 �� 57 ��� 24 ������E, R/V Cape St. Mary, cr. 7 / 64, st. 32, Agassiz trawl, 823���834 m, 5 Nov. 1964. Nontypes: USNM 207354, 2 females (1 cleared and stained) and ZMUC P 77550, female, 95 mm SL, off Hawaiian Islands, 21 ��06���N, 156 �� 13 ���W, R/V Townsend Cromwell, 52���88, shrimp trawl, 585���640 m, 3 Mar. 1971. BSKU 29526, female, 138 mm SL, Kyusu-Palau ridge, 28 ��04���N, 134 �� 41 ���E, R/V Shinkai-Maru, No. 53, beam trawl, 523 m, 18 Dec. 1979. AMS I. 22814 -029, female, 160 mm SL, off Northwest Australia, 18 �� 29 ���S, 116 �� 36 ���E, R/V Soela, st. SO 2 / 82 / 26,27, Engel trawl, 696���704 m, 6 Apr. 1982. AMS I. 26002 -005, female, 84 mm SL, off New South Wales, 33 �� 34 ���S, 152 �� 2 ���E, R/V Kapala, st. K 86 -01-06, bottom trawl, 658 m, 11 Feb. 1986. IOM 0 0 498, male, 94 mm SL, Mozambique Channel, 12 �� 28 ��� 30 ������S, 48 ��09��� 54 ������E to 12 �� 26 ���S, 48 ��08 E, R/V Vityaz, cr. 17, st. 2601, bottom trawl, 700���710 m, 12 Nov. 1988. AMS I. 29749 -002, female, 90 mm SL, off New South Wales, 32 �� 40 ���S, 152 �� 51 ���E to 32 �� 38 ���S, 152 �� 54 ���E, R/V Kapala, K 89 - 17 -02, bottom trawl, 814���850 m, 15 Aug. 1989. MNHN 2002-3143, female, 119 mm SL, Chesterfield Islands, 19.7 ��S, 158.83 ��E, Musorstom 5, st. cc 384, bottom trawl, 756���772 m, 21 Oct. 1986. MNHN 2000 -0677, female, 147 mm SL, Vanua Balavu, Fiji Islands, 17.33 ��S, 178.7 ��W, R/V Buge, cr. 40, st. cp 1431, bottom trawl, 495���500 m, 2 Mar. 1999. MNHN 2006 -0588, female, 100 mm SL, Solomon Islands, 8.38 ��S, 159.47 ��E, Camp. Salomon 2, st. cp 2194, bottom trawl, 440���521 m, 24 Oct. 2004. MNHN 2006 -0702, male, 112 mm SL, Solomon Islands, 7.62 ��S, 157.75 ��E, Camp. Solomon 2, cr. cp 2214, 550 ��� 682 m, 26 Oct. 2004. MNHN 2006 -0090, male, 113 mm SL, Solomon Islands, 7.71 �� S, 156.47 ��E, Camp. Salomon 2, st. cp 2246, bottom trawl, 664���682 m, 1 Nov. 2004. MNHN 2006 -0464, male, 84 mm SL, Solomon Islands, 7.7 ��S, 156.43 ��E, Camp. Salomon 2, st. cp 2247, bottom trawl, 686���690 m, 1 Nov. 2004. MNHN 2006 -0061, 2 females, 91���98 mm SL, Solomon Islands, 7.9 ��S, 156.95 ��E, Camp. Salomon 2, st. cp 2267, bottom trawl, 590���600 m, 4 Nov. 2004. IOM (no cat. no.), 137 mm SL, off Vietnam, R/V Odissey tr. 52; no further data available. Diagnosis. Saccogaster tuberculata differs from the other Saccogaster species by the following combination of characters: No body scales; a pair of small, sub-dermal spines on frontal plate just behind eyes, median subdermal ethmoidal spine in front of eyes; skin rather thick; large posterior mandibular pore behind end of maxilla; cartilaginous ethmoidal spine on snout; opercular spine hidden, no preopercular spine; three developed rakers on anterior gill arch almost equal in length to developed rakers; pectoral peduncle prolonged; no fangs on dentigerous bones, palatines with 3 to several tooth rows; precaudal vertebrae 14���15 and total vertebrae 55���58; fin rays in dorsal 82���93, anal 54���60 and pectoral 19���22; otolith length to height 2���2.1, otolith length to colliculum length variable, 2.5���3.3. Similarity. Saccogaster tuberculata seems most close to S. horrida, both having developed gill rakers on anterior arch equal to or longer than long filaments, three developed gill rakers and no scales. They differ from each other by S. tuberculata having an indistinct frontal plate and small subdermal spines (vs. a distinct frontal plate with a pair of well developed, visible spines), no transverse, bony ridge on frontal plate (vs. bony ridge present), presence of a posterior mandibular pore (vs. absent), dentigerous bones without fangs (vs. vomer and premaxillaries with fangs), distance from base of pelvic fins to origin of anal fin 32.0���39.0 % SL (vs. 46.0 % SL) and ratio otolith length to height 2.1���2.2 (vs. 1.75). Description. The principal meristic and morphometric characters are shown in Table 6. Body compressed and elongate with tapering caudal part and blunt snout. Scales absent. Skin relatively thick, loose and semitransparent. Lateral line indistinct, originating above gill opening, descends gradually above anus and continues in midline becoming obscure posteriorly. In some specimens the 25���30 lateral line papillae are black clearly showing the course of the line. Dorsal fin origin above vertebrae 6���8 and anal fin origin below vertebrae 21���23. Pectoral fin reaching more than halfway to anus. Anterior gill arch (Fig. 18 F) with 3���4 spiny knobs on upper branch, a developed raker in angle and lower branch with two developed rakers with a knob between followed by 7���11 knobs. Number of knobs varies between 14 and 19. Two pseudobranchial filaments. Intromittent organ of holotype on a thick, fleshy, posteriorly directed stalk with the vent immediately anterior to its base; distal rim of stalk swollen and divided into two lobes, with a small penis in the middle; sections of the testes show spermatozoans arranged in spermatophores. In USNM 207354 (SL 130 mm) ovaries contain several hundred dark-eyed, uniformly brownish embryos with yolk sacs. S. staigeri S. tuberculata Axial skeleton (based on radiographs): Number of precaudal vertebrae 14���15. Anterior neural spine one fourth length of second spine. Neural spines 2 to 4 ��� 5 long and compressed and with pointed tips, spines 6���11 with blunt tips and vertebrae 4���8 with enlarged basal part. Parapophyses on vertebrae 7 to 14 ��� 15, pleural ribs on vertebrae 2���10 and epipleural ribs on vertebrae 4���10. Pleural and epipleural ribs often indistinct. Dentition: Fangs absent. Dentition granular. Palatines with 3 to several, irregular tooth rows. Vomer boomerang-shaped and with 3���4 tooth rows. Premaxillaries and dentaries with several, irregular tooth rows. Number of tooth rows seems to increase with standard length. Head morphology (Fig. 18 A���E): Head profile straight above eyes. Head with pair of small, sub-dermal spines on frontal plate above and behind eyes, small, median, sub-dermal, ethmoidal spine in front of eyes and blunt subdermal spine above eyes. Sub-dermal surface of frontal plate behind frontal spines slightly textured. Anterior nostril placed close to upper lip; posterior nostril a mere hole placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, loose, tighter in small specimens and then translucent and sometimes with faint indications of 1���3 subdermal neuromasts below and behind eyes. No head pigmentation visible. Upper jaw ends well behind eye; posterior end of maxilla vertically expanded. Opercular spine pointed, subdermal, not reaching hind margin of opercle. No spine at lower angle of preoperculum. Otolith (Fig. 18 G���J): Otolith elongate, moderately thick; otolith length to height = 2.0��� 2.2, otolith height to thickness = 1.9���2.2. Ventral rim regularly curved, smooth; dorsal rim regularly curved or broadly undulating. Anterior tip moderately pointed; posterior tip slightly pointed or rounded. Inner face flat with centrally positioned, undivided oval, shallow sulcus with distinct anterior projection and somewhat elevated colliculum; otolith length to colliculum length variable, 2.5���3.3. Outer face markedly convex. Coloration: Recently caught specimens often with brownish snout, dark dorsum and black papillae on head and in lateral line. Biology and distribution (Fig. 25). The 18 examined specimens were caught in bottom fishing gear at depths of 440���850 m from off East Africa to Hawaii. Remarks on variability. Saccogaster tuberculata is the most widely distributed species of the genus. It also shows a remarkably high variation of certain characters, which however is not consistent enough to warrant distinction of separate species. Particularly it appears that it is always single characters that make the distinction of any given variation, which is not supported by any independent second character. Two specimens from Hawaii, the northeastern-most occurrence of the species, show otoliths with a regularly curved dorsal rim (Fig. 18 G), whereas all others exhibit an undulating dorsal rim. The southwestern-most specimen, from off Madagascar, has a tight head skin (vs. mostly loose) and a distinctly narrower interorbital width (3.4 % SL vs. 3.9���5.6 % SL). A specimen from off New South Wales, Australia, has a single, feeble neuromast (Fig. 18 E) and two specimens from off the Solomon Islands show 2 and 3 neuromasts, respectively, all on the upper lip behind the eye. A few specimens from off the Solomon Islands show dusky pigmentation on the posterior part of the body., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 22-26, DOI: 10.5281/zenodo.208677, {"references":["Chan, W. (1966) A new genus and species of deep-sea brotulid from the South China Sea. Japanese Journal of Ichthyology, 1 (3), 4 - 8.","Cohen, D. M. (1987) Notes on the bythitid fish genus Saccogaster with a new species from the Gulf of Mexico. Contribution in Science, 385, 1 - 4","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp."]}

Saccogaster maculata Alcock, 1889 Figs. 9, 10, 25, Table 1, 2, 5 Saccogaster maculatus Alcock 1889: 389 (type locality: Bay of Bengal). Saccogaster maculata: Alcock 1891: 226, fig.; Wood-Mason & Alcock 1891: 30, pl. 7, fig. 3.; Alcock & McArdle 1900: pl. 29, figs. 2 and 2 a (intromittent organ); Cohen & Nielsen 1972: 448, fig. 2; Cohen 1987: 1; Nielsen et al. 1999: 109. Saccogaster maculatus: Menon & Yazdani 1968: 150 (types in ZSI). Diplacanthopoma (Saccogaster) maculatum: Alcock 1899: 102. Material examined (5 specimens and radiograph of syntype (ZSI 11673), 75���87 mm SL). ZSI 13527, female, 81 mm SL and ZSI 13528, female, 75 mm SL, Bay of Bengal, R/V Investigator, bottom trawl, 265��� 458 m. ZSI 13045, male, 77 mm SL, Bay of Bengal, R/V Investigator, bottom trawl, 439 m. IOM no cat. no., male, 87 mm SL, Bay of Bengal, 16 �� 56 ���08������N, 83 �� 13 ���04������E, R/V Vityaz, cr. 33, st. 4929, bottom trawl, 600 m, 1 Jan. 1961. IOM, no cat. no., 72 mm SL, Gulf of Aden, 10 �� 32 ��� 12 ������N, 49 �� 22 ��� 12 ������E, R/V Dmitry Stefanov, cr. 5, st. 98, bottom trawl, 450 ��� 445 m, 6 Dec. 1989. Remarks on material: It was not possible to borrow any of the five specimens from ZSI for this revision and the IOM specimens seem lost so information about S. maculata is based on Cohen & Nielsen (1972: 448���451). Consequently, our information on ethmoid and frontal spines is not based on direct observation, but from available radiographs. Diagnosis. Saccogaster maculata differs from all other Saccogaster species by the following combination of characters: Distinct pair of (probably) free spines on frontal plate above and behind eyes, strong ethmoidal spine in front of eyes; scales present on sides of body but not on predorsal; some specimens with ventral arm of opercle terminating in a short spine; opercular spine free and with one prong; presence of 2 posterior infraorbital pores, 1 supraorbital pore at upper angle of gill opening and 1 posterior mandibular behind maxillary; anterior gill arch with three long rakers about half length of longest filaments; elongate pectoral peduncle; palatines with several tooth rows; precaudal vertebrae 12���13 and total vertebrae 53��� 55; fin rays in dorsal 83���85, anal 53���56 and pectoral 17���18. Similarity. Saccogaster maculata is most similar to S. staigeri as both are scaled, have gill filaments 1���2 times length of developed rakers and fangs on dentigerous bones. They differ by S. maculata having palatines with several tooth rows (vs. single tooth row) and opercular spine free (vs. hidden). Also S. maculata has posterior infraorbital, supraorbital and posterior mandibular pores, which are all missing in S. staigeri. Description ( from Cohen & Nielsen 1972: 449) The principal meristic and morphometric characters are shown in Table 5. Head and body elongate and compressed with tapering tail and blunt snout. Skin thin and transparent. Scales absent on predorsal and in a band along bases of dorsal and anal fins except posteriorly, where entire body is scaled. Anteriorly scales widely scattered, posteriorly scales imbricate. The scale pattern in Alcock and McArdle (1900, pl. 29, fig. 2) is inaccurate. Lateral line (Fig. 9) in two non-overlapping sections; anterior, dorsal part with about 12 small papillae extending from upper angle of opercle to about anus; posterior, median part with 8���10 small papillae ending somewhat anterior to base of caudal fin. Dorsal fin origin above pectoral peduncle and anal fin origin well behind midpoint of fish. Pectoral fin extending to more than half the distance from upper angle of peduncle to anal fin; peduncle longer than broad and free of body. Snout depressed and rounded. Small, darkly pigmented papillae and dermal fringes sparsely distributed on snout and a few on posterior part of head. Posterior tip of opercle ends in blunt spine. In some specimens ventral arm of opercle ends in a short spine. Upper jaw ends well behind eyes with posterior part vertically expanded. Anterior gill arch (Fig. 10 B) with 3���4 spiny knobs on upper branch, one developed raker in angle and lower branch with two developed rakers with a knob in between followed by 12 spiny knobs. Gill filaments 2���3 times length of developed rakers. Two pseudobranchial filaments. Intromittent organ a thick, fleshy, posteriorly-directed stalk with two prominent lobes on tip from which a penis protrudes. Paired testes of ���Vityaz��� specimen not notably enlarged; a histological section shows spermatozoa arranged in spermatophores. Axial skeleton (from radiographs): Number of precaudal vertebrae 13. Anterior neural spine considerably shorter than second spine. Neural spines on precaudal vertebrae 2���5 with pointed tips and decreasing in length, spines 6���9 with blunt tips and remaining neural and haemal spines with pointed tips. Parapophyses on vertebrae 7���13. Ribs difficult to observe. Dentition: Palatines with sharp, pointed teeth in a narrow band several teeth wide. Vomer boomerang-shaped with sharp, pointed teeth. Premaxillaries with band of granular teeth, but near symphysis an inner row of 7���9 elongate, needlelike teeth. Dentaries with a band of granular teeth and an inner row of about 15 pointed teeth. Head pores (from Cohen & Nielsen 1972: 449): 1 supraorbital pore at tip of snout and 1 at upper angle of gill opening, 3 anterior infraorbital pores below and in front of eye and 2 posterior infraorbital pores below and behind eyes, 2���3 anterior mandibular pore at tip of jaw, 1 posterior mandibular pore behind posterior end of maxillary. Head morphology from radiograph (Fig. 10 A): Pair of forward directed strong spines, probably visible, on frontal above and behind eyes, strong upward directed median ethmoidal spine in front of eyes and blunt subdermal spine above eyes. The surface of the frontal plate behind the frontal spines appears to be roughly textured. Otolith: No data available. Coloration: Except for dusky pigmentation on snout preserved specimens are without color patterns. Biology and distribution (Fig. 25). A viviparous species caught in bottom fishing gear on the upper continental slope at depths between 265 and 600 m. Six specimens were caught in the Gulf of Bengal and one in the Gulf of Aden. *Dorsal fin, anal fin and vertebral counts from only two specimens, Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 14-16, DOI: 10.5281/zenodo.208677, {"references":["Alcock, A. (1889) Natural history notes from H. M. Indian marine survey steamer \" Investigator \". - No. 13. On the bathybial fishes of the Bay of Bengal and neighbouring waters during the seasons 1885 - 1889. Annals and Magazine of Natural History, ser. 6, 4, 376 - 399.","Wood-Mason, J. & Alcock, J. (1891) Natural history notes from H. M. Indian marine survey steamer \" Investigator \". Ser. II, No. 1. On the result of the deep-sea dredging during the seasons 1890 - 91. Annals and Magazine of Natural History, ser. 6, 8, 16 - 34, pl. 7, fig. 3.","Alcock, A. & McArdle, A. F. (1900) Illustrations of the zoology of the royal Indian marine survey ship \" Investigator \". Fishes - Part 7, Royal Indian Marine, Calcutta, pls. 27 - 35.","Cohen, D. M. & Nielsen, J. G. (1972) A review of the viviparous ophidioid fishes of the genus Saccogaster. Proceedings of the Biological Society of Washington, 85 (39), 445 - 468.","Cohen, D. M. (1987) Notes on the bythitid fish genus Saccogaster with a new species from the Gulf of Mexico. Contribution in Science, 385, 1 - 4","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp.","Menon, A. G. K. & Yazdani, G. M. (1968) Catalogue of type-specimens in the Zoological Survey of India, Part. 2. Fishes. Records of the Zoological Survey of India, 61 (1 - 2), 91 - 190.","Alcock, A. (1899) A Descriptive Catalogue of the Indian Deep-Sea Fishes in the Indian Museum. Indian Museum, Calcutta. Pp. 1 - 211."]}

Saccogaster brayae n. sp. Figs. 3, 4, 25, Table 1, 2, 4 Material examined (1 specimen, 36 mm, SL). Holotype: NMV A 29655 -002, male, 36 mm SL, off northwestern Australia, Barrow L 1 transect, 21 ��00��� 49 ������S, 114 �� 39 ���09������E to 21 ��00��� 47 ������S, 114 �� 38 ��� 54 ������E, R/V Southern Surveyor, Sherman sledge, 250���258 m, 10 June 2007. Diagnosis. Saccogaster brayae differs from all other Saccogaster species in having a broken brown stripe between eye and hind margin of opercle (except for S. hawaii), precaudal vertebrae 21 and total vertebrae 53, and becoming ripe at a very small SL (36 mm). Also the following combination of characters is diagnostic: A pair of free spines on frontal above and behind eyes, a large median sub-dermal ethmoidal spine in front of eyes with forward pointed tip; antero-ventrally directed spine at lower angle of preopercle; thin transparent skin; scales present on sides of body; three long gill rakers on anterior arch 4���5 times the length of gill filaments; palatines with 1���2 rows of teeth; pectoral peduncle longer than high; fin rays in dorsal 85, anal 52 and pectoral 18; otolith length to height = 1.9, otolith length to colliculum length = 3.1, sulcus without anterior projection. Similarity. Saccogaster brayae seems most similar to S. hawaii by the brown stripe behind eye, the forward pointed tip of the ethmoidal spine and the scaled body. They differ by S. brayae having opercular spine ending in one prong (vs. three prongs), by long rakers on anterior arch 4���5 times length of filaments (vs. long rakers and filaments almost equal in length), precaudal vertebrae 21 (vs. 18), pectoral fin rays 18 (vs. 22) and dorsal fin rays 85 (vs. 92). Description. The principal meristic and morphometric characters are shown in Table 4. Head and body elongate and compressed with tapering caudal part and blunt snout. Most scales rubbed off but scale-pockets and a few remaining scales (0.8 mm long) indicate squamation on sides of body. Lateral line not visible. Dorsal fin origin above proximal part of pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fin ending less than halfway to anus with peduncle a little longer than broad and free of body. Anterior gill arch with three spiny knobs on upper branch, a developed raker in angle and lower branch with two developed rakers with a knob in between followed by eight knobs. Developed rakers 4���5 times length of gill filaments (Fig. 4 D). Two small pseudobranchial filaments. The well developed testes and intromittent organ (Fig. 4 G) show that the specimen is a ripe male. Axial skeleton (from radiographs): Number of precaudal vertebrae 21. Anterior neural spine very short. Neural spines 2���4 long, depressed and with pointed tips. Neural spines 5���20 short and with large basal parts, length of spines increasing posteriorad. Parapophyses developed on vertebrae 6���20. Pleural ribs distinct at vertebrae 1���8, but pleural ribs on remaining precaudal vertebrae and presence of epipleural ribs are questionable. Dentition: Palatines with about 25 pointed teeth in 1���2 rows; larger teeth in inner row. Vomer boomerangshaped with about 25 pointed teeth in 2���3 irregular rows; six longer teeth in inner row. Premaxillaries with granular teeth except for five fangs at symphysis. Dentaries with 3���4 irregular tooth rows, outer rows with granular teeth and inner row with fangs. Head morphology (Fig. 4 A���C): Head profile concave above eyes. Head with pair of free spines as an integrated part of a transverse rim on frontal above and slightly behind eyes and a large, median sub-dermal ethmoidal spine in front of eyes with forward pointed tip. Anterior nostril placed close to upper lip and small posterior nostril placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, tight. No neuromasts visible. Head with dusky background pigmentation, broken brown stripe between eye and hind margin of opercle and broken brown pigmentation patches below and behind eye. Upper jaw ends well behind eye with posterior margin vertically expanded. Opercular spine pointed, just reaching hind margin of operculum. Antero-ventrally directed spine at lower angle of preoperculum. Otolith (Fig. 4 E���F): Otolith oval, moderately thick; otolith length to height = 1.9, otolith height to thickness = 1.9. Dorsal and ventral rims gently curved, dorsal rim broadly undulating. Anterior and posterior tips regularly rounded resulting in a regular oval appearance of the otolith. Inner face flat with centrally positioned, undivided, small, oval, shallow sulcus without anterior projection. Otolith length to colliculum length = 3.1. Outer face convex. Coloration: After five years of preservation body uniformly brownish with faint dark pigmentation dorsally. Head with dusky, dense pigmentation and a dark brown stripe from eye to hind margin of opercle. Larger, dark pigment spots on frontal plate seen through skin on dorsum. Eyes blue with yellow lens. Biology and distribution (Fig. 25). A viviparous species occurring on the upper continental slope. Known from a 36 mm male with well developed testes and intromittent organ, caught off northwestern Australia organ (Fig. 4 G), caught at a depth of 250��� 258 m. Etymology. Named after Dianne Bray, MNV, for her support with material for this revision., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 7-9, DOI: 10.5281/zenodo.208677

Key to Hastatobythites, Saccogaster and Parasaccogaster n. gen. 1 Gill opening ends at or below level of dorsal margin of pectoral peduncle; pectoral peduncle attached to body; skin thick; eye sunk in, subdermal............................................................. Parasaccogaster n. gen. (p. 26) - Gill opening ends above level of dorsal margin of pectoral peduncle; pectoral peduncle free of body; skin thin (except in S. tuberculata); eye not subdermal.......................................................................... 2 2 Narrow head (5.6 ���6.0 % SL) and body (3.5���3.9 % SL); one anteriorly directed median spine above eyes; three bony ridges on frontal plate......................................................................... Hastatobythites (p. 3) - Broad head (10.5 ���15.0 % SL) and body (4.2���9.5 % SL); pair of spines above and behind eyes (absent in S. parva); no or single bony ridge on frontal plate................................................................ Saccogaster (p. 5), Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 2-3, DOI: 10.5281/zenodo.208677

Hastatobythites Machida, 1997 Hastatobythites Machida 1997: 385, type species by original designation Hastatobythites arafurensis Machida, 1997. Hastatobythites: Nielsen et al. 1999: 105. Diagnosis. A monotypic genus of the subfamily Bythitinae (Cohen & Nielsen 1978: 42) with the following characters: two spines placed in midline of head, one anteriorly directed on frontal above eye and another upward directed, weak and thin on ethmoid, three bony ridges on dorsum behind frontal spine; maximum width of head 5.6 ���6.0% SL and of body 3.5���3.9 % SL. Also the following combination of characters is diagnostic: Elongate body with joined vertical fins; head naked and anterior part of body with scattered scales in midline and posterior half of body almost fully scaled; skin thin, translucent; eye diameter less than snout-length; opercular spine covered by skin; posterior part of maxilla greatly expanded vertically; dentigerous bones with granular teeth; palatines with 2���3 tooth rows; pectoral peduncle slightly longer than broad, not adnate; gill opening extending well above opercular spine; anterior gill arch with three long rakers; precaudal vertebrae 15 and total vertebrae 57���58; fin rays in dorsal 99���100, caudal 12, anal 64���65 and pectoral 16���17; otolith length to height = 2.5, sulcus undivided, placed in central part of inner face, otolith length to sulcus length = 2.7. Similarity. Hastatobythites is most similar to Saccogaster in having more or less prolonged pectoral radials, naked head, teeth on palatines and thin skin. Hastatobythites differs from Saccogaster by having one median spine on frontal plate (vs. a pair or none), very narrow head (5.6 ���6.0 % SL vs. 10.5 ���15.0 % SL) and body (3.5���3.9 % SL vs. 4.2���9.5 % SL) and three median bony ridges behind frontal spine (absent or one in Saccogaster). Hastatobythites differs from Parasaccogaster n. gen. in the very narrow head, three median bony ridges behind frontal spine, prolonged free pectoral radials (vs. adnate) and the thin, tight head skin (vs. thick and loose). Pectoral Dorsal Anal Species 75 77 79 81 83 85 87 89 91 93 95 97 99 37 39 41 43 45 47 49 51 53 55 57 59 61 63 65 12 13 14 15 16 17 18 19 20 21 22 23 ��� 74 ��� 76 78 ��� 80 ��� 82 ��� 84 ��� ��� 86 ��� 88 ��� 90 92 ��� ��� 94 96 ��� 98 ��� 100 ��� 36 ��� 38 ��� 40 42 ��� ��� 44 46��� 48 ��� 50 ��� ��� 52 54��� 56 ��� 58 ��� 60 ��� 62 ��� 64 ��� H. arafurensis + + + + + P. melanomycter + + + P. normae + + + + + + + + P. rhamphidognatha + + + S. brayi + + + S. hawaii + + + S. horrida + + + S. maculata + + + + + + + S. nikoliviae + + + + + + + + S. parva + + + S. staigeri + + + + + + + S. tuberculata + + + + + + + + + + + + + +, Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 3-4, DOI: 10.5281/zenodo.208677, {"references":["Machida, Y. (1997) Hastatobythites arafurensis, a new bythitid genus and species from Australia (Bythitidae, Ohidiiformes). Ichthyological Research, 44, 385 - 388.","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp.","Cohen, D. M. & Nielsen, J. G. (1978) Guide to the identification of genera of the fish order Ophidiiformes with a tentative classification of the order. NOAA Technical Report NMFS Circular, 417, 1 - 72."]}

Hastatobythites arafurensis Machida, 1997 Figs. 1, 2, 25, Tables 1���3 Hastatobythites arafurensis Machida 1997: 386, figs. 1���2 (type locality: Arafura Sea, 9 �� 18 ���S, 133 �� 25 ���E). Material examined (1 specimen and photo and radiograph of holotype NTM S. 12904 -011): Paratype: NTM S. 12904 -018, female, 57 mm SL, NNE of Money Shoal, Arafura Sea. 9 �� 18 ���S, 133 �� 25 ���E, otter trawl, 146 m, 7 Nov. 1990. Remarks on material: Two specimens known from the same locality. A specimen (AMS I. 29749 -002) caught off New South Wales was originally identified as H. arafurensis and cited by Nielsen et al. (1999: 105). It is now correctly referred to Saccogaster tuberculata (Chan, 1966). Diagnosis. See diagnosis for Hastatobythites Description. The major meristic and morphometric characters are shown in Table 3. Head and body elongate and strongly compressed with tapering caudal part and blunt snout. Posterior part of body fully scaled, anterior part with a few scales in midline and head naked. No visible lateral line. Dorsal fin origin above proximal part of pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fin ending less than halfway to anus with peduncle a little longer than broad. Anterior gill arch (Fig. 2 D) with 4���5 knobs on upper branch, a developed raker in angle and lower branch with two developed rakers with a knob between followed by 11 knobs; all rakers and knobs with small, densely placed spines. Long rakers about twice the length of gill filaments. Two small pseudobranchial filaments. Head pores on paratype difficult to observe. Axial skeleton (based on radiographs): Number of precaudal vertebrae 15. Anterior neural spine one fifth the length of second spine. Neural spines 4���5 to 6���7 gradually decreasing in length. Pointed tips on neural spines 1���2 to 4���5, blunt tips on spines 6 to 9 ��� 11 and remaining vertebrae with neural and haemal spines pointed. Vertebrae 3���4 to 6 with depressed tips and vertebrae 3���7 with enlarged basal parts. Parapophyses developed on vertebrae 6���15. Pleural and epipleural ribs show up so poorly on radiographs that counts are questionable. Dentition (based on paratype): Vomer and palatines with granular dentition in irregular rows and no fangs. Dentaries with 15���20 fangs in inner row and many small, pointed teeth in irregular outer rows. Premaxillaries with many small, pointed teeth in irregular rows and a few fangs near symphysis. Head morphology (Fig. 2 A���C): Head profile with minor concavity above eyes. Two spines in midline of head; a larger, free, forward-directed frontal spine placed above eyes and a smaller, ethmoidal spine partly covered by skin in front of eyes. Three median bony ridges on dorsum behind frontal spine (length 6.5 % SL). Anterior nostril placed close to upper lip and larger posterior nostril placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, tight. No neuromasts visible. No head pigmentation visible. Upper jaw ends well behind eye with posterior margin vertically expanded. Opercular spine flat but pointed not reaching hind margin of opercle. No spine at lower angle of preoperculum. Otolith (Fig. 2 E���F): Otolith elongate, thin; otolith length to height = 2.5, otolith height to thickness = 2.1. Dorsal and ventral rims regularly and shallow curved, dorsal rim with slight predorsal lobe. Anterior and posterior tips regularly rounded resulting in an oval appearance of the otolith. Inner face flat with centrally positioned, undivided small, oval, shallow sulcus; otolith length to colliculum length = 2.7. Outer face flat. Coloration. Paratype uniformly lightbrown, dorsal part of head darker and eyes bluish with light lens. Biology and distribution (Fig. 25). A viviparous species occurring on the lower continental shelf. Holotype a male with simple intromittant organ. Paratype a female with 4 embryos (4���5 mm long) with red-brown eyes, up to 20 scattered, red-brown spots on body and a short, brown stripe above abdomen (Fig. 1 C). Known from two specimens caught in the same trawl haul in the Arafura Sea from a depth of 146 m., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 4-7, DOI: 10.5281/zenodo.208677, {"references":["Machida, Y. (1997) Hastatobythites arafurensis, a new bythitid genus and species from Australia (Bythitidae, Ohidiiformes). Ichthyological Research, 44, 385 - 388.","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp.","Chan, W. (1966) A new genus and species of deep-sea brotulid from the South China Sea. Japanese Journal of Ichthyology, 1 (3), 4 - 8."]}

Saccogaster staigeri Cohen & Nielsen, 1972 Figs. 15, 16, 26, Table 1, 2, 6 Saccogaster staigeri Cohen & Nielsen 1972: 2, fig. 1 (type locality: east coast of Florida). Saccogaster staigeri: Cohen 1987: 2; Nielsen et al. 1999: 109. ? Saccogaster staigeri: S��ret & Andreatta 1992: 92 (off Brazil). Material examined (5 specimens, 49���93 mm SL)*. Holotype: USNM 207357, male, 93 mm SL, east coast of Florida, 27 �� 11 ���N, 79 �� 49 ���W, R/V Gerda, st. 657, 10��� otter trawl, 201���216 m, 16 July 1965. Paratype: USNM 164144, female, 61 mm SL, off Tortugas, Gulf of Mexico, 24 �� 20 ��� N, 82 �� 55 ���W, R/V Oregon, st. 1005, 40��� shrimp trawl, 347 m, bottom temp. 10.0��C, 13 Apr. 1954. Non-types: USNM 158640, female, 72 mm SL, off Dry Tortugas, Florida, 24 �� 20 ���N, 83 �� 20 ���W, R/V Oregon, st. 1005, 347 m, 13 Apr. 1954. TCWC 6146.1, female, 65 mm SL, northwestern Gulf of Mexico, 27 �� 44.9 ���N, 95 �� 20.1 ���W, R/V Alaminos, st. 68 A 13 ���19, 2 m dredge, 338���384 m, 19 Nov. 1968. TCWC 7027.20, male, 49 mm SL, northern Gulf of Mexico, 28 �� 28 ��� 47 ������N, 86 �� 2 ��� 32 ������W, R/V Gyre, st. E1, 9 m otter trawl, 351���357 m, 14 May 1985. * The following specimen of S. staigeri was reported upon by S��ret and Andreatta (1992) but it has not been available to us for checking the identification. It is tentatively referred to S. staigeri: USU 0 1501, 80 mm SL, off Rio de Janeiro, 23 �� 41.64 ���S, 42 ��06.53���W, R/V Marion Dufresne, st. 63 CB 104, Blake trawl, 430���450 m, 1 June, 1987. Diagnosis. Saccogaster staigeri differs from all other Saccogaster species by the following combination of characters: Frontal plate with a pair of small, hidden, sub-dermal spines above and behind eyes and a small, median, sub-dermal, ethmoidal spine in front of eyes; opercular spine hidden by skin, preopercular spine absent; pectoral peduncle prolonged; three developed rakers on anterior gill arch 1.5���2 times length of gill filaments; palatines with one tooth row and fang-like teeth on all dentigerous bones; precaudal vertebrae 13���14 and total vertebrae 53���54; fin rays dorsal 86���88, in anal 53���57 and pectoral 18���19; otolith length to height = 2.1, otolith length to colliculum length = 2.8. Similarity. Saccogaster staigeri is most similar to S. maculata as both are scaled, have gill filaments 1.5���2 times length of developed rakers and fangs on dentigerous bones. They differ by S. staigeri having palatines with a single tooth row (vs. several tooth rows) and opercular spine and pair of spines on frontals hidden (vs. free). Also S. staigeri has no posterior infraorbital, supraorbital and posterior mandibular pores, which all are present in S. maculata. Description. The principal meristic and morphometric characters are shown in Table 6. Body moderately compressed and elongate with tapering caudal part and blunt snout. Scales on body concentrated around midline, hardly overlapping; no scales on head and abdomen. Lower part of lateral line in holotype with about 20 papillae. Dorsal fin origin above proximal part of pectorals and anal fin origin well behind midpoint of fish. Pectoral fin ending almost halfway to anus with prolonged peduncle not attached to body. Pelvic fins ending below anterior part of pectoral fins. Anterior gill arch (Fig. 16 D) with four flat, spiny knobs on upper branch, a developed raker in the angle and lower branch with two longer rakes with a knob in between followed by 11 low knobs. Developed rakers on anterior arch 1.5���2 times length of filaments. Two pseudobranchial filaments. Holotype with 9 mm long penis and rear part of body cavity with swollen testes. Eggs are visible through the body wall of paratype. Axial skeleton (from radiographs): Number of precaudal vertebrae 13���14. Anterior neural spine very short. Neural spines 2���4 long and with pointed tips and neural spines 5���12 distinctly shorter, with blunt tips and enlarged basis. All remaining neural and haemal spines with pointed tips. Parapophyses developed on vertebrae 7 to 13 ��� 14. Pleural ribs on vertebrae 3���4 to 9. Epipleural ribs on vertebrae 3���9. Dentition: Vomer boomerang-shaped with 7���8 fangs in 1���2 rows. Palatines with 10���12 fangs in a single row. Dentaries with 15���20 fangs in one row. Premaxillaries with granular teeth in many rows and 2���3 fangs near symphysis. Head morphology (Fig. 16 A���C): Head profile straight above eyes. Head with pair of tiny sub-dermal spines on frontal plate above and behind eyes, distinct on radiographs, small, median, sub-dermal, ethomoidal spine in front of eyes and blunt sub-dermal spine above eyes. Frontal plate behind frontal spines with coarsely textured subdermal surface. Anterior nostril placed close to upper lip; posterior nostril a mere hole placed close to eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thin, loose. No neuromasts visible. No head pigmentation visible. Upper jaw ends well behind eye; posterior end of maxilla vertically expanded. Opercular spine pointed, subdermal, not reaching hind margin of opercle. No spine at lower angle of preoperculum. Otolith (Fig. 16 E���F): Otolith elongate, moderately thick; otolith length to height = 2.1, otolith height to thickness = 2.3. Dorsal and ventral rims regularly curved, smooth. Anterior and posterior tips pointed resulting in a symmetrical outline of the otolith. Inner face slightly convex with centrally positioned, undivided, oval, shallow sulcus without anterior projection; otolith length to colliculum length = 2.8. Outer face convex. Coloration: After about 50 years of preservation the specimens are uniformly brownish except for the darkblue eyes. Biology and distribution (Fig. 25). Caught in bottom fishing gear at the upper continental slope at a depth of 201��� 450 m. The holotype is from off the east coast of Florida and the paratype and three more specimens from the Gulf of Mexico. According to S��ret and Andreatta (1992) a fifth specimen was trawled off Rio de Janeiro, which is here recorded as tentatively assigned., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 20-22, DOI: 10.5281/zenodo.208677, {"references":["Cohen, D. M. & Nielsen, J. G. (1972) A review of the viviparous ophidioid fishes of the genus Saccogaster. Proceedings of the Biological Society of Washington, 85 (39), 445 - 468.","Cohen, D. M. (1987) Notes on the bythitid fish genus Saccogaster with a new species from the Gulf of Mexico. Contribution in Science, 385, 1 - 4","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp.","Seret, B. & Andreata, J. V. (1992) Deep-sea fishes collected during cruise MD- 55 off Brazil. Cybium, 16 (1), 81 - 100."]}

Key to the species of Saccogaster 1 Scales present on body................................................................................ 2 - Scales absent from body................................................................................ 6 2 Opercular spine free, with 3 prongs; pectoral fin rays 22; dorsal fin rays 92.................................. hawaii - Opercular spine free or hidden, with single prong; pectoral fin rays 16���19; dorsal fin rays 80���88....................... 3 3 Gill filaments on anterior arch short (1 / 3 ��� 1 / 5 times length of developed rakers)................................... 4 - Gill filaments on anterior arch about equal to or longer than developed gill rakers.................................. 5 4 Scales present on predorsal; 12���13 precaudal vertebrae; no brown stripe on head; numerous neuromasts...... nikoliviae n.sp. - Scales absent on predorsal; 21 precaudal vertebrae; broken brown stripe on sides of head; no neuromasts....... brayae n.sp. 5 Gill filaments on anterior arch distinctly longer than developed gill rakers; opercular spine free; palatines with several tooth rows; dorsal fin rays 83���85; posterior infraorbital pores 2, posterior supraorbital pore 1, posterior mandibular pore 1....... maculata - Gill filaments on anterior arch 0.5���1 times length of developed gill rakers; opercular spine hidden; palatines with a single tooth row; dorsal fin rays 86���88; no posterior infraorbital pores, no posterior supraorbital pore, no posterior mandibular pore..................................................................................................... staigeri 6 Developed gill rakers on anterior arch 5���6; pectoral fin rays 14; numerous neuromasts........................... parva - Developed gill rakers on anterior arch 3; pectoral fin rays 19���22; 0���3 neuromasts................................... 7 7 Skin thin; opercular spine free; spines on frontal plate visible, bony ridges above eyes; distance from base of pelvic fins to ori- gin of anal fin 46.0% SL; vomer and premaxillaries with fangs; no posterior mandibular pore.................... horrida - Skin thick; opercular spine hidden; spines on frontal plate sub-dermal, no bony ridge above eyes; distance from base of pelvic fins to origin of anal fin 32.0���39.0 % SL; no fangs, dentition granular; large posterior mandibular pore......... tuberculata, Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on page 7, DOI: 10.5281/zenodo.208677

Parasaccogaster normae (Cohen & Nielsen, 1972) Figs. 21, 22, 26, Tables 1, 2, 7 Saccogaster normae Cohen & Nielsen 1972: 463, figs. 1 and 6 (type locality: off northern Peru). Saccogaster normae: Cohen 1987: 1; Nielsen et al. 1999: 110. Material examined (6 specimens, 118–142 mm SL). Holotype: USNM 207356, female, 140 mm SL, off northern Peru, 4 ° 57 ’S, 81 ° 23 ’W, R/V Anton Bruun, cr. 16, st. 625 A, otter trawl, 118–133 m, 2 June 1966. Paratype: IM 421, male, 138 mm SL, off northern Peru, 5 °04’S, 80 ° 24 ’W, R/V Lance, st. 76, 150 m, 7 May 1969. Non-types: UCR 1555 - 5 (male, 102 mm SL), UCR 1555 - 8 (female, 128 mm SL) and UCR 2072 - 21 (female, 118 mm SL), Peninsula de Nicoya, Costa Rica, 9 ° 44 ’N, 85 ° 22 ’W, R/V Nautilus III, bottom trawl, 30 Oct. 1983. UCR 760 - 1, female, 142 mm SL, off Quepos, Puntarenos Prov., Costa Rica, 9 ° 24 ’ 50 ’’N, 85 ° 22 ’W, bottom trawl, 25 Mar. 1973. Diagnosis. Parasaccogaster normae differs from the other two species of the genus by the gill filaments on anterior arch being 4–5 times the length of the developed rakers and by the following combination of characters: A pair of very small, subdermal, closely set spines on frontal plate above and behind eyes, no spine on ethmoid; 2–3 developed rakers on anterior arch not much longer than spiny knobs; gill filaments on anterior arch 4–5 times length of developed rakers; palatines with 2–4 tooth rows; pectoral peduncle prolonged and adnate to body; precaudal vertebrae 14–16 and total vertebrae 54–56; fin rays in dorsal 83–87, caudal 12–13, anal 51–54 and pectoral 17–19. Similarity. Parasaccogaster normae is most similar to P. rhamphidognatha with hidden opercular spine, nasal chamber not black and distinctly more anal fin rays and vertebrae than found in S. melanomycter. Parasaccogaster normae differs from P. rhamphidognatha by the longer gill filaments on anterior arch (4–5 times length of long rakers vs. 1–3 times), more pectoral fin rays (17–19 vs. 12) and lack of an ethmoidal spine vs. spine present. Description. The principal meristic and morphometric characters are shown in Table 7. Head and body elongate with tapering caudal part and blunt snout. Scales absent. Origin of lateral line dorsally above pectoral peduncle and from above anus running in midline ending at base of caudal fin with 27 to ca. 40 small, dark papillae. Dorsal fin origin above anterior part of pectoral fin and anal fin well behind midpoint of fish. Pectoral fin ending far from halfway to anal fin and with prolonged peduncle. Anterior gill arch (Fig. 22 E) with 3–4 small, spiny knobs on upper branch, one a little longer raker in the angle and lower branch with two short rakers with a knob in between followed by ca. ten spiny knobs. Long gill filaments 4–5 times length of developed rakers. Two pseudobranchial filaments. Ovaries large, but no embryos observed. Axial skeleton (from radiographs): Number of precaudal vertebrae 15–16. Anterior neural spine one fourth length of second spine. Neural spines 2–4 long and depressed and with pointed tips, spines 5–10 with blunt tips and nos. 5–8 depressed and with enlarged basal parts. Parapophyses developed on vertebrae 8–15, pleural ribs on vertebrae 3–10 and epipleural ribs not observed. Dentition: Palatines with 2–3 rows of pointed teeth, largest in inner row. Vomer boomerang-shaped and with 2–3 irregular rows of pointed teeth, largest in inner row. Premaxillaries with several irregular rows of pointed teeth and a few fangs at symphysis. Dentaries with pointed teeth in irregular rows, largest in inner row. Head morphology (Fig. 22 A–B): Head profile slightly or not concave above eyes. Head with pair of very small, sub-dermal, closely set spines on frontal plate above and behind eyes, no spine on ethmoid, broad, blunt, sub-dermal spine above eyes, frontal plate with a median, bony ridge. Anterior nostril placed close to upper lip and larger posterior nostril placed close to small eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thick, loose. Head with blunt snout, sunk-in eye covered by window of translucent skin. Upper jaw ends well behind eye with posterior margin vertically expanded. Opercular spine hidden below thick skin. No subdermal preopercular spine. Gill opening ending slightly below level of dorsal margin of pectoral peduncle. Otolith (Fig. 22 C–D): Otolith elongate, thin; otolith length to height = 2.3, otolith height to thickness = 2.5. Dorsal and ventral rims gently and regularly curved, smooth. Anterior and posterior tips pointed resulting in a symmetrical outline of the otolith. Inner face slightly convex with centrally positioned, undivided oval, shallow sulcus without anterior projection; otolith length to colliculum length = 2.5. Outer face nearly flat. Coloration: Head and body chocolate brown with darker fins. Biology and distribution (Fig. 26). A viviparous species living on the lower part of the continental shelf (80–150 m). Four of the examined specimens are females with well developed eggs but apparently no embryos. Sections of ripe testes show that the spermatozoans are arranged in spermatophores (Cohen and Nielsen 1972: 465). Known from off northern Peru and from the Pacific coasts of Costa Rica.

Parasaccogaster melanomycter (Cohen, 1981) Figs. 19, 20, 26, Table 1, 2, 7 Saccogaster melanomycter Cohen 1981: 374, figs. 1���2 (type locality: Gairaca Bay, Caribbean Columbia). Saccogaster melanomycter: Cohen 1987: 2; Nielsen et al. 1999: 110. Material examined (1 specimen, 66 mm SL). Holotype: MCZ 47362, 66 mm SL, female, Santa Maria, Gairaca Bay (Encenada de Gayraca), Caribbean Columbia, coral reef, 8 m, 1968. Diagnosis. Parasaccogaster melanomycter differs from the other two Parasaccogaster species by the sooty black nasal chambers, few anal fin rays (37 vs. 49���54), few vertebrae (45 vs. 54���58) and a wider head (12.0 % SL vs. 7.7���9.8 % SL). Also the following combination of characters is diagnostic: A pair of sub-dermal spines fused to form a broad ridge on frontal just behind eyes, a small, median, sub-dermal, ethmoidal spine just in front of eyes and an antero-ventrally directed spine at lower angle of preopercle (all spines covered by skin); three developed rakers on anterior gill arch only slightly larger than the spiny knobs, gill filaments 7���8 times length of developed rakers; pectoral peduncle slightly prolonged; palatines with several tooth rows; precaudal vertebrae 16 and total vertebrae 45; fin rays in dorsal 74, anal 37, caudal 10 and pectoral 18. Similarity. Parasaccogaster melanomycter differs from the two other species of the genus in the stouter body shape, fewer vertebrae (45 vs 54���58), fewer anal fin rays (37 vs 49���54) and wider head (12.0 % SL vs 7.7���9.8 % SL). Also the fusion of the paired frontal spines is not observed in any other species of the genera studied here. Description. The principal meristic and morphometric characters are shown in Table 7. Body rather compressed with pointed, slightly compressed head and tapering tail. Skin thick and opaque. Scales absent. Lateral line continuous, originating above and anterior to upper angle of gill opening and descending to midline beyond midpoint of body. Its course marked by narrow unpigmented line with widely spaced, small papillae; about 12 dark papillae to level of vent and further posteriorly pale papillae difficult to count. Dorsal fin origin above middle of pectoral fin and anal fin origin well behind midpoint of fish. Pectoral fin reaching 1 / 3 to anal fin; pectoral peduncle longer than broad and adnate to body. Anterior gill arch (Fig. 20 D) with five low, spiny knobs on upper branch, one slightly developed raker in angle and lower branch with two slightly developed rakers with a low knob in between and followed by five partly united knobs, two shallow, likewise partly united knobs, and four well developed separate knobs after a small break. Longest gill filaments about 2���3 three times length of developed rakers. Two pseudobranchial filaments. Body cavity nearly filled with two distended ovaries packed with apparently unfertilized eggs of about 1 mm in diameter. Axial skeleton (from radiograph): Number of precaudal vertebrae 16. Anterior neural spine very short and nos. 2���4 very long. Neural spines 2���8 much depressed and with blunt tips. Parapophyses developed on vertebrae 6���16. Pleural ribs on vertebrae 3���11. Dentition: Palatines with rows of pointed teeth, inner row with larger teeth. Vomer with a boomerang-shaped band of sharp-pointed teeth, inner row with larger teeth. Premaxillaries with a narrow band of granular teeth and an inner row of enlarged, needle-like teeth. Dentaries with irregular row of small, slightly compressed teeth and an inner row of larger, pointed teeth. Head morphology (Fig. 20 A���C, E���F): A pair of fused, sub-dermal spines on frontal plate above and just behind eyes, a small, thin, compressed, sub-dermal, ethmoidal spine in midline of snout and broad, blunt, subdermal spine above eyes. Frontal plate with rough sub-dermal texture. Opercular spine pointed, covered by skin, not reaching hind margin of operculum. Antero-ventrally directed spine at lower angle of preopercle below skin. Anterior nostril placed close to upper lip, not tubular; posterior nostril twice as large as anterior nostril, positioned at some distance from eye. Head pores: 1 supraorbital pore at tip of snout, 1 small posterior supraorbital pore above base of opercular spine, 2 anterior infraorbital pores below and in front of eye, 2 anterior mandibular pores at tip of jaw. Head with pointed snout, sunk-in eye covered by window of translucent skin, and many small, dark papillae on occiput and cheeks. Upper jaw ends well behind eye with posterior part vertically expanded. Nasal chambers sooty black about equal in size to orbit. Otolith: Dissolved. Coloration: Color light brown, darker dorsally and on caudal section. Dorsal and anal fins with narrow, pale rim. Nasal chamber lined with sooty, black pigment, separated from eye by white area. Head papillae darker than skin color. Biology and distribution (Fig. 26). Parasaccogaster melanomycter is only known from the holotype caught at a depth of 8 m on a coral reef in Gairaca Bay, Caribbean Columbia. It is the only species of the three genera here reviewed living on a shallow reef., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 27-29, DOI: 10.5281/zenodo.208677, {"references":["Cohen, D. M. (1981) Saccogaster melanomycter (Ophidiiformes: Bythitidae), a new fish species from the Caribbean. Proceedings from the Biological Society of Washington, 94 (2), 374 - 377.","Cohen, D. M. (1987) Notes on the bythitid fish genus Saccogaster with a new species from the Gulf of Mexico. Contribution in Science, 385, 1 - 4","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp."]}

Parasaccogaster normae (Cohen & Nielsen, 1972) Figs. 21, 22, 26, Tables 1, 2, 7 Saccogaster normae Cohen & Nielsen 1972: 463, figs. 1 and 6 (type locality: off northern Peru). Saccogaster normae: Cohen 1987: 1; Nielsen et al. 1999: 110. Material examined (6 specimens, 118���142 mm SL). Holotype: USNM 207356, female, 140 mm SL, off northern Peru, 4 �� 57 ���S, 81 �� 23 ���W, R/V Anton Bruun, cr. 16, st. 625 A, otter trawl, 118���133 m, 2 June 1966. Paratype: IM 421, male, 138 mm SL, off northern Peru, 5 ��04���S, 80 �� 24 ���W, R/V Lance, st. 76, 150 m, 7 May 1969. Non-types: UCR 1555 - 5 (male, 102 mm SL), UCR 1555 - 8 (female, 128 mm SL) and UCR 2072 - 21 (female, 118 mm SL), Peninsula de Nicoya, Costa Rica, 9 �� 44 ���N, 85 �� 22 ���W, R/V Nautilus III, bottom trawl, 30 Oct. 1983. UCR 760 - 1, female, 142 mm SL, off Quepos, Puntarenos Prov., Costa Rica, 9 �� 24 ��� 50 ������N, 85 �� 22 ���W, bottom trawl, 25 Mar. 1973. Diagnosis. Parasaccogaster normae differs from the other two species of the genus by the gill filaments on anterior arch being 4���5 times the length of the developed rakers and by the following combination of characters: A pair of very small, subdermal, closely set spines on frontal plate above and behind eyes, no spine on ethmoid; 2���3 developed rakers on anterior arch not much longer than spiny knobs; gill filaments on anterior arch 4���5 times length of developed rakers; palatines with 2���4 tooth rows; pectoral peduncle prolonged and adnate to body; precaudal vertebrae 14���16 and total vertebrae 54���56; fin rays in dorsal 83���87, caudal 12���13, anal 51���54 and pectoral 17���19. Similarity. Parasaccogaster normae is most similar to P. rhamphidognatha with hidden opercular spine, nasal chamber not black and distinctly more anal fin rays and vertebrae than found in S. melanomycter. Parasaccogaster normae differs from P. rhamphidognatha by the longer gill filaments on anterior arch (4���5 times length of long rakers vs. 1���3 times), more pectoral fin rays (17���19 vs. 12) and lack of an ethmoidal spine vs. spine present. Description. The principal meristic and morphometric characters are shown in Table 7. Head and body elongate with tapering caudal part and blunt snout. Scales absent. Origin of lateral line dorsally above pectoral peduncle and from above anus running in midline ending at base of caudal fin with 27 to ca. 40 small, dark papillae. Dorsal fin origin above anterior part of pectoral fin and anal fin well behind midpoint of fish. Pectoral fin ending far from halfway to anal fin and with prolonged peduncle. Anterior gill arch (Fig. 22 E) with 3���4 small, spiny knobs on upper branch, one a little longer raker in the angle and lower branch with two short rakers with a knob in between followed by ca. ten spiny knobs. Long gill filaments 4���5 times length of developed rakers. Two pseudobranchial filaments. Ovaries large, but no embryos observed. Axial skeleton (from radiographs): Number of precaudal vertebrae 15���16. Anterior neural spine one fourth length of second spine. Neural spines 2���4 long and depressed and with pointed tips, spines 5���10 with blunt tips and nos. 5���8 depressed and with enlarged basal parts. Parapophyses developed on vertebrae 8���15, pleural ribs on vertebrae 3���10 and epipleural ribs not observed. Dentition: Palatines with 2���3 rows of pointed teeth, largest in inner row. Vomer boomerang-shaped and with 2���3 irregular rows of pointed teeth, largest in inner row. Premaxillaries with several irregular rows of pointed teeth and a few fangs at symphysis. Dentaries with pointed teeth in irregular rows, largest in inner row. Head morphology (Fig. 22 A���B): Head profile slightly or not concave above eyes. Head with pair of very small, sub-dermal, closely set spines on frontal plate above and behind eyes, no spine on ethmoid, broad, blunt, sub-dermal spine above eyes, frontal plate with a median, bony ridge. Anterior nostril placed close to upper lip and larger posterior nostril placed close to small eye. Head pores: 1 supraorbital pore at tip of snout, 2 anterior infraorbital pores below and in front of eye, 1 anterior mandibular pore at tip of jaw. Head skin thick, loose. Head with blunt snout, sunk-in eye covered by window of translucent skin. Upper jaw ends well behind eye with posterior margin vertically expanded. Opercular spine hidden below thick skin. No subdermal preopercular spine. Gill opening ending slightly below level of dorsal margin of pectoral peduncle. Otolith (Fig. 22 C���D): Otolith elongate, thin; otolith length to height = 2.3, otolith height to thickness = 2.5. Dorsal and ventral rims gently and regularly curved, smooth. Anterior and posterior tips pointed resulting in a symmetrical outline of the otolith. Inner face slightly convex with centrally positioned, undivided oval, shallow sulcus without anterior projection; otolith length to colliculum length = 2.5. Outer face nearly flat. Coloration: Head and body chocolate brown with darker fins. Biology and distribution (Fig. 26). A viviparous species living on the lower part of the continental shelf (80���150 m). Four of the examined specimens are females with well developed eggs but apparently no embryos. Sections of ripe testes show that the spermatozoans are arranged in spermatophores (Cohen and Nielsen 1972: 465). Known from off northern Peru and from the Pacific coasts of Costa Rica., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on pages 29-31, DOI: 10.5281/zenodo.208677, {"references":["Cohen, D. M. & Nielsen, J. G. (1972) A review of the viviparous ophidioid fishes of the genus Saccogaster. Proceedings of the Biological Society of Washington, 85 (39), 445 - 468.","Cohen, D. M. (1987) Notes on the bythitid fish genus Saccogaster with a new species from the Gulf of Mexico. Contribution in Science, 385, 1 - 4","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp."]}

Saccogaster Alcock, 1889 Saccogaster Alcock 1889: 389, type species by monotypy Saccogaster maculata Alcock, 1889. Barbuliceps Chan 1966: 4, type species by original designation Barbuliceps tubercularis (lapsus pro tuberculatus) Chan, 1966. Saccogaster: Cohen & Nielsen 1972; Nielsen et al. 1999: 109. Diagnosis. A genus of the subfamily Bythitinae (Cohen & Nielsen 1972) characterized by the following combination of characters: A pair of bony spines free or sub-dermal on frontal plate above and behind eyes and one median spine on ethmoid (except none in S. parva); elongate body with joined vertical fins; anal fin origin behind midpoint of fish; pectoral fins with elongated radials, not adnate; skin thin, translucent (except thick in adult S. tuberculata); scales absent on head and reduced or absent on body; opercular spine present, free in some species, covered by skin in others; maxilla vertically expanded posteriorly; palatine teeth present; gill opening extending well above dorsal margin of pectoral peduncle; developed gill rakers on anterior arch 3���6; pseudobranchial filaments 2; fin rays in dorsal 80���93, anal 51���64 and pectoral fin 14���22; precaudal vertebrae 12���21, total vertebrae 51���58; males with stalked intromittent organ. Similarity. The most similar genus is Hastatobythites (see p. 3). Species. The genus Saccogaster contains eight species known from 41 specimens of which three seem to be lost and two are only accessible at ZSI, Kolkata. Four are known from one specimen only, three species from 3��� 6 specimens and one species from 18 specimens., Published as part of Nielsen, J��rgen G., Schwarzhans, Werner & Cohen, Daniel M., 2012, Revision of Hastatobythites and Saccogaster (Teleostei, Bythitidae) with three new species and a new genus, pp. 1-36 in Zootaxa 3579 on page 7, DOI: 10.5281/zenodo.208677, {"references":["Alcock, A. (1889) Natural history notes from H. M. Indian marine survey steamer \" Investigator \". - No. 13. On the bathybial fishes of the Bay of Bengal and neighbouring waters during the seasons 1885 - 1889. Annals and Magazine of Natural History, ser. 6, 4, 376 - 399.","Chan, W. (1966) A new genus and species of deep-sea brotulid from the South China Sea. Japanese Journal of Ichthyology, 1 (3), 4 - 8.","Cohen, D. M. & Nielsen, J. G. (1972) A review of the viviparous ophidioid fishes of the genus Saccogaster. Proceedings of the Biological Society of Washington, 85 (39), 445 - 468.","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk-eels, brotulas and other ophidiiform fishes known to date. FAO Species Catalogue, 18. FAO Fish. Synopsis, 125 (18), 178 pp."]}

Bellottia cryptica, new species (Figs. 4���7) Bellottia apoda: Ross & Quattrini (2007: 985, 996) Bellottia n. sp.: Ross & Quattrini (2008: 80) Material examined (1 specimen, SL 42 mm): Holotype: NCSM 44451, SL 42, female, off southern Georgia, 30 �� 48.74 ���N, 79 �� 37.96 ���W, st. JSL-04- 4485, 629 m, coll. S.W. Ross, 11 June 2004. Condition of material. The holotype and only specimen was eviscerated for ecological studies, but is otherwise in good condition. Organs removed are archived with the specimen. Diagnosis. Bellottia cryptica differs from its congeners by the following combination of characters: body slender (17.0 % SL at origin of anal fin), teeth small and pointed, caudal fin rays 7, origin of anal fin below dorsal fin ray 8, anterior gill arch with 5 long rakers, predorsal length 46.5 % SL, precaudal vertebrae 10, cleithrum with distinct spine just above base of pectoral fin, no spine on preopercular crest. Similarity. Bellottia cryptica is most similar to B. robusta, in e. g. presence of spine on cleithrum and in fin ray and total vertebral counts, but differs by the more slender body (17.0 vs 17.5 ���23.0 % SL at origin of anal fin), origin of anal fin below dorsal fin ray 8 vs 10���15, precaudal vertebrae 10 vs 12 and no spine on preopercular crest vs 1 spine in B. robusta. It differs from B. galatheae by having 5���6 long rakers on anterior gill arch vs 4, and by having more rays in dorsal (92 vs 70���72), anal (84 vs 61) and pectoral (23 vs 19���20) fins. From B. apoda it differs by having 10 precaudal vertebrae vs 12, spine on cleithrum vs spine absent and 5���6 long rakers on anterior gill arch vs 3���4. From B. armiger it differs by having 7 caudal fin rays (vs 6), total vertebrae 49 (vs 45) and no spine on preopercular crest (vs spine present). Description. The principal meristic and morphometric characters are shown in Table 1. Body short, highest near base of pectoral fin, completely covered with oval ca. 0.7 mm cycloid scales. Lateral line indistinct with anterior part apparently near dorsal margin of body. Head profile convex, cheek and gill cover scaled with rest of head naked. Mouth oblique with posterior part of maxilla vertically expanded ending just behind eye. Anterior nostril with low rim placed closer to upper lip than to posterior nostril. Small opercular spine strong and pointed, reaching beyond posterior margin of opercle. Lower edge of preopercle with four distinct spines, and no spine on preopercular crest (Fig. 3). Origin of dorsal fin above tip of pectoral fin, anal fin origin at midpoint of fish. Pectoral fin placed on mid-body level with peduncle higher than long. A prominent spine on cleithrum just above base of pectoral fin. Anterior gill arch with 3���4 platelike rakers on upper branch, one long raker in the angle between the two branches and lower branch with 4���5 long rakers followed by 8���9 plate-like rakers. Longest gill filament about 1 / 3 of longest raker. Two short, broad pseudobranchial filaments. Head pores: The loose and partly torn skin makes it difficult to observe the head pores. The following large, distinct pores were seen: three behind and three below the eye and four pores on the mandible. A few whitish sensory papillae distributed over the head. Dentition: Head of vomer slightly curved with about 10 very small, blunt teeth in one row. Posterior half of palatines edentate, anterior half with small teeth in one row. Posterior half of dentaries edentate, anterior half with one row increasing to 4���5 tooth rows at symphysis with large and pointed teeth in outer row. Premaxillaries with posterior fourth edentate and further anteriorly small teeth in one row increasing to 4���5 rows at symphysis with larger and pointed teeth in outer row. Axial skeleton (based on radiographs): Tips of neural and haemal spines thin and pointed. Anterior neural spine one third the length of second spine. Neural spines 2���8 depressed. Bases of neural spines 6���8 enlarged. Parapophyses present on vertebrae 7���10 and pleural ribs on vertebrae 4���9. Epipleural ribs indistinct. Otolith (Fig. 6): The 2 mm elongate sagittal otolith is 1.5 times as long as high and twice as high as thick. Dorsal and ventral rim rounded, vaguely pointed anteriorly. Sulcus undivided and 1 / 3 ��� 1 / 4 the length of the otolith. No osteal channel. Coloration: Fig. 5 shows the holotype just after capture. The body is light brown, the margin of the vertical fins dark, the abdomen dark blue, the gill cover and side of head reddish brown and the eye is dark brown. Biology. The general benthic habitat in the area where this specimen was collected consisted of a rugged, rocky ledge that was colonized by scattered hard corals (mostly Lophelia pertusa), black corals, bamboo corals, sponges and hydroids. Bottom temperature was 7.8 �� C and salinity was 34.9 ppt. This specimen was swimming about 1.5���2 m above the ledge and was collected by suction sampler. A large amount of rotenone was spread in the area, and it seems likely this fish was escaping from that. We assume that this species is normally benthic and cryptic, similar to B. robusta. The stomach of this specimen was about one third full of unidentified crustacean parts. Etymology. The specific name, cryptica, refers to its hidden nature. Distribution (Fig. 4). Known from a single specimen caught off southern Georgia, 30 �� 48.74 ���N, 79 �� 37.96 ���W, by suction-sampling from JSL at a depth of 629 m. We predict that it has a wider distribution along the slope between North Carolina and southeastern Florida., Published as part of Nielsen, J��rgen G., Ross, Steve W. & Cohen, Daniel M., 2009, Atlantic occurrence of the genus Bellottia (Teleostei, Bythitidae) with two new species from the Western North Atlantic, pp. 45-57 in Zootaxa 2018 on pages 51-53, DOI: 10.5281/zenodo.186025, {"references":["Ross, S. W. & Quattrini, A. M. (2008) Deep - sea reef fish assemblage patterns on the Blake Plateau. Marine Ecology, 30, 74 - 92."]}

Bellottia cryptica, new species (Figs. 4–7) Bellottia apoda: Ross & Quattrini (2007: 985, 996) Bellottia n. sp.: Ross & Quattrini (2008: 80) Material examined (1 specimen, SL 42 mm): Holotype: NCSM 44451, SL 42, female, off southern Georgia, 30 ° 48.74 ’N, 79 ° 37.96 ’W, st. JSL-04- 4485, 629 m, coll. S.W. Ross, 11 June 2004. Condition of material. The holotype and only specimen was eviscerated for ecological studies, but is otherwise in good condition. Organs removed are archived with the specimen. Diagnosis. Bellottia cryptica differs from its congeners by the following combination of characters: body slender (17.0 % SL at origin of anal fin), teeth small and pointed, caudal fin rays 7, origin of anal fin below dorsal fin ray 8, anterior gill arch with 5 long rakers, predorsal length 46.5 % SL, precaudal vertebrae 10, cleithrum with distinct spine just above base of pectoral fin, no spine on preopercular crest. Similarity. Bellottia cryptica is most similar to B. robusta, in e. g. presence of spine on cleithrum and in fin ray and total vertebral counts, but differs by the more slender body (17.0 vs 17.5 –23.0 % SL at origin of anal fin), origin of anal fin below dorsal fin ray 8 vs 10–15, precaudal vertebrae 10 vs 12 and no spine on preopercular crest vs 1 spine in B. robusta. It differs from B. galatheae by having 5–6 long rakers on anterior gill arch vs 4, and by having more rays in dorsal (92 vs 70–72), anal (84 vs 61) and pectoral (23 vs 19–20) fins. From B. apoda it differs by having 10 precaudal vertebrae vs 12, spine on cleithrum vs spine absent and 5–6 long rakers on anterior gill arch vs 3–4. From B. armiger it differs by having 7 caudal fin rays (vs 6), total vertebrae 49 (vs 45) and no spine on preopercular crest (vs spine present). Description. The principal meristic and morphometric characters are shown in Table 1. Body short, highest near base of pectoral fin, completely covered with oval ca. 0.7 mm cycloid scales. Lateral line indistinct with anterior part apparently near dorsal margin of body. Head profile convex, cheek and gill cover scaled with rest of head naked. Mouth oblique with posterior part of maxilla vertically expanded ending just behind eye. Anterior nostril with low rim placed closer to upper lip than to posterior nostril. Small opercular spine strong and pointed, reaching beyond posterior margin of opercle. Lower edge of preopercle with four distinct spines, and no spine on preopercular crest (Fig. 3). Origin of dorsal fin above tip of pectoral fin, anal fin origin at midpoint of fish. Pectoral fin placed on mid-body level with peduncle higher than long. A prominent spine on cleithrum just above base of pectoral fin. Anterior gill arch with 3–4 platelike rakers on upper branch, one long raker in the angle between the two branches and lower branch with 4–5 long rakers followed by 8–9 plate-like rakers. Longest gill filament about 1 / 3 of longest raker. Two short, broad pseudobranchial filaments. Head pores: The loose and partly torn skin makes it difficult to observe the head pores. The following large, distinct pores were seen: three behind and three below the eye and four pores on the mandible. A few whitish sensory papillae distributed over the head. Dentition: Head of vomer slightly curved with about 10 very small, blunt teeth in one row. Posterior half of palatines edentate, anterior half with small teeth in one row. Posterior half of dentaries edentate, anterior half with one row increasing to 4–5 tooth rows at symphysis with large and pointed teeth in outer row. Premaxillaries with posterior fourth edentate and further anteriorly small teeth in one row increasing to 4–5 rows at symphysis with larger and pointed teeth in outer row. Axial skeleton (based on radiographs): Tips of neural and haemal spines thin and pointed. Anterior neural spine one third the length of second spine. Neural spines 2–8 depressed. Bases of neural spines 6–8 enlarged. Parapophyses present on vertebrae 7–10 and pleural ribs on vertebrae 4–9. Epipleural ribs indistinct. Otolith (Fig. 6): The 2 mm elongate sagittal otolith is 1.5 times as long as high and twice as high as thick. Dorsal and ventral rim rounded, vaguely pointed anteriorly. Sulcus undivided and 1 / 3 – 1 / 4 the length of the otolith. No osteal channel. Coloration: Fig. 5 shows the holotype just after capture. The body is light brown, the margin of the vertical fins dark, the abdomen dark blue, the gill cover and side of head reddish brown and the eye is dark brown. Biology. The general benthic habitat in the area where this specimen was collected consisted of a rugged, rocky ledge that was colonized by scattered hard corals (mostly Lophelia pertusa), black corals, bamboo corals, sponges and hydroids. Bottom temperature was 7.8 ° C and salinity was 34.9 ppt. This specimen was swimming about 1.5–2 m above the ledge and was collected by suction sampler. A large amount of rotenone was spread in the area, and it seems likely this fish was escaping from that. We assume that this species is normally benthic and cryptic, similar to B. robusta. The stomach of this specimen was about one third full of unidentified crustacean parts. Etymology. The specific name, cryptica, refers to its hidden nature. Distribution (Fig. 4). Known from a single specimen caught off southern Georgia, 30 ° 48.74 ’N, 79 ° 37.96 ’W, by suction-sampling from JSL at a depth of 629 m. We predict that it has a wider distribution along the slope between North Carolina and southeastern Florida.

Key to species of Bellottia spp. 1 a Long rakers on anterior gill arch 5–6............................................................................................................................ 2 1 b Long rakers on anterior gill arch 3–4............................................................................................................................ 4 2 a Caudal fin rays 6, precaudal vertebrae 10, total vertebrae 45, anal fin rays 74 ............ B. armiger (Philippine Islands) 2 b Caudal fin rays 7, precaudal vertebrae 10–12, total vertebrae 48–51, anal fin rays 75–88.......................................... 3 3 a Precaudal vertebrae 12, preopercular crest with spine (Fig. 3), depth at origin of anal fin 17.5 –23.0 % of SL........... ........................................................................................................................................... B. robusta (Gulf of Mexico) 3 b Precaudal vertebrae 10, preopercular crest without spine, depth at origin of anal fin 17.0 % SL................................. .................................................................................................................................. B. cryptica (off southern Georgia) 4 a Dorsal fin rays 70–72, pectoral fin rays 19–20, origin of anal fin below dorsal fin ray no. 12–13, predorsal 44.0– 45.5 % SL, spine on cleithrum present, no fangs on dentigerous bones................................................................................ ...................................................................................................... B. galatheae (Solomon Sea and Philippine Islands) 4 b Dorsal fin rays 83–94, pectoral fin rays 21–26, origin of anal fin below dorsal fin ray no. 17–19, predorsal 35.5 –40.0 % SL, spine on cleithrum absent, all dentigerous bones with retrorse fangs................................................................. .................................................................................................................... B. apoda (Mediterranean and NE Atlantic)

Bellottia apoda Giglioli, 1883 (Figs. 8���10) Bellottia apoda Giglioli, 1883: 399 (type locality Gulf of Naples, Italy); Nielsen & Cohen 1968: 99 (selection and illustration of lectotype MZUF 5668); Nielsen et al. 1968: 245 (spermatophores); Maul 1976: 53 (off Portugal and on Great Meteor Bank southwest of Madeira); Orsi 1976: 38 (Ligurian Sea); Papaconstantinou 1984: 103 (Gulf of Evoikos, Greece); Berdar et al. 1977: 77 (Strait of Messina); Papaconstantinou et al. 1977: 279 (Gulf of Saronikos, Greece); Jardas 1979: 1 (Adriatic Sea); Sardou 1980: 69 (Villefranche-sur-Mer, South France; illustration of head pore system); Schwarzhans 1981: 103 (otolith); Allu�� 1984: 41 (near Barcelona); de Ranieri & Sbrana 1992: 177 (Northern Tyrrhenian Sea); Gramito & Coen 1997: 163 (Adriatic Sea); Anon. 2002 (Mallorca) - non Bellottia apoda; Mytilineou et al. 2005: 230 (eastern Ionian Sea). Material examined (16 specimens, SL 26���65 mm): Lectotype: MZUF 5668, SL 28, male, Gulf of Naples, trawl, Posidonia- bottom, 30 m, 20 Dec. 1882. - Paralectotype: MZUF 5669, SL 26, female, same data as for lectotype. Non-type material: MMF 22516, SL 52���55, male and female, off Portugal, 37 �� 41.5 ���N, 9 �� 11.9 ���W, R/V Meteor, st. 11 a-AT 7, Agassiz trawl, 500 m, 20 Jan. 1967. MMF. 22860, SL 39 + - 65,?- male, off Portugal, 37 �� 21.5 ���N, 9 �� 12.5 ���W, R/V Meteor, st. 90 d-AT 26, Agassiz trawl, 320���385 m, 22 June 1967. ZMUC P 77698, SL 46, male, Great Meteor Bank, 29 �� 50.2 ���N, 28 �� 29.8 ���W, R/V Meteor, st. 159 a, bottom trawl, 308���310 m, 18 July 1967. ZMUC P 771651, SL 38, female, Central Adriatic Sea, 42 �� 51 ���N, 14 �� 44.6 ���E, bottom trawl, 247 m, 30 Jan. 1980. BMNH 1999.3. 23.1, SL 41, male, Adriatic Sea, 42 �� 48.5 ���N, 14 ��48.0���E, 18 June 1980. IIPB 454 / 1982 and 455 / 1982, SL 47���52, male and female, off Barcelona, 41 �� 6.05 ���N, 2 �� 3.1 ���E, bottom trawl, 446���516 m, 30 June 1981. ZMUC P 771646 - 1650, SL 29���45, males, Central Adriatic Sea, 42 �� 48.9 ���N, 14 �� 41.5 ���E, bottom trawl, 217���219 m, 15 Apr. 1994. Diagnosis. Bellottia apoda differs from all its congeners by having retrorse fangs on all dentigerous bones and lacking a distinct spine on cleithrum above base of pectoral fin and by the following combination of characters: body slender (13.0���16.0 % SL at origin of anal fin), spine on preopercular crest, caudal fin rays 6, anterior gill arch with 3���4 long rakers, precaudal vertebrae 12���13 and predorsal length (35.5 ���40.0). Similarity. Judging from Table 2, B. apoda seems to differ equally much from all the other Bellottia species making it difficult to select one species as the most similar. It differs from B. robusta e.g. by a more slender (13.0���16.0 vs 17.5 ���23.0 % SL) and more light-colored body, by fewer caudal fin rays (6 vs 7), fewer long rakers on anterior gill arch (3���4 vs 5���6), lacking spine on cleithrum (spine present) and shorter predorsal length (35.5 ���40.0 vs 44.0��� 49.5). From B. cryptica it differs e.g. by having fewer caudal fin rays (6 vs 7), more precaudal vertebrae (12���13 vs 10), fewer long rakers on anterior gill arch (3���4 vs 5���6) and presence of a spine on preopercular crest (spine absent). From B. armiger it differs e.g. by having fewer long rakers on anterior gill arch (3���4 vs 5), more precaudal vertebrae (12���13 vs 10), and origin of anal fin below dorsal fin ray 17���19 (vs 11). From B. galatheae it differs i. a. by having more rays in dorsal fin (83���94 vs 70���72), anal fin (68���82 vs 61), pectoral fin (21���26 vs 19���20), and a shorter predorsal length (35.5 ���40.0 vs 44.0��� 45.5). Description. The principal meristic and morphometric characters are shown in Table 1. Nielsen & M��ller (2008, table 3) used the same set of characters as in the present paper based on 13 specimens three of which they examined while the characters of the rest were taken from the literature. For comparative reasons we have here only included characters from the 16 specimens examined by us. B. apoda is thoroughly described and illustrated by Nielsen & Cohen (1968), Sardou (1980) and Gramitto & Coen (1997), rendering a detailed description here redundant. A few characters that have not been focused on in earlier papers are mentioned below. Dentition. In males vomer, palatines and dentaries with many large, retrorse fangs; premaxillaries with small, pointed teeth besides fangs. In females fangs on all four dentigerous bones, but fewer and shorter than in males. apoda armiger galatheae robusta cryptica Axial skeleton (based on radiographs): Tips of neural and haemal spines thin and pointed. Anterior neural spine 1 / 2 to 1 / 1 the length of second spine. Neural spines 2���9 depressed. Bases of neural spines 4���9 enlarged. Parapophyses present on vertebrae 7 / 8 to 12 / 13, pleural ribs on vertebrae 2 / 3 to 10 / 11 and epipleural ribs on vertebrae 4 / 5 to 11 / 12; the latter observed on few specimens only. Otolith (Fig. 9). The 3.9 mm elongate sagittal otolith (from a 65 mm SL specimen, MMF 22860) is twice as long as high and twice as high as thick. Dorsal and ventral rim evenly rounded. Sulcus undivided, almost half as long as the otolith. No osteal channel. Coloration: Fig. 8 shows a specimen after 27 years of preservation. It is uniformly light brown except for the dark eyes. A photo (provided by E. Gramitto, Ancona) of a specimen of unknown length from the Adriatic Sea taken just after capture shows a light brown body with darker vertical fins, a dark brown head with many spots and dark blue eyes. Biology. All specimens have been caught on or near the bottom. The type material was trawled at 30 m on Posidonia -bottom and all specimens since then at depths of 150��� 569 m. This is a viviparous genus in which the male transfers the spermatozoans packed in spermatophores into the female (Nielsen et al. 1968: 245). A 38 mm specimen (ZMUC P 771651) contained eggs up to 1 mm in diameter; embryos were not observed. Gramitto & Coen (1997: 170) observed embryos ready to hatch of 5 mm total length. According to Gramitto & Coen (1997: 168), the stomach contents from 23 Adriatic specimens showed that the diet consisted of pelagic and benthic crustaceans with copepods as the most important prey. Distribution. Fig. 10 shows that B. apoda is found from Greece to off Barcelona, off southern Portugal and furthermore on Great Meteor Bank at depths of 30��� 569 m. Remarks. B. apoda seems to be a rather common species in the Mediterranean as 128 specimens were caught in the Central Adriatic Sea between 1970 and 1994 (Gramitto & Coen 1997: 163). In order to determine whether the specimens from the East Atlantic Ocean differ from the Mediterranean ones, we have compared the set of characters used in Table 1 between these areas. Overlap was observed in all but one character, viz. number of pectoral fin rays: 21���23 in the Atlantic and 23���26 in the Mediterranean specimens, suggesting that there is no difference between the two populations., Published as part of Nielsen, J��rgen G., Ross, Steve W. & Cohen, Daniel M., 2009, Atlantic occurrence of the genus Bellottia (Teleostei, Bythitidae) with two new species from the Western North Atlantic, pp. 45-57 in Zootaxa 2018 on pages 53-56, DOI: 10.5281/zenodo.186025, {"references":["Giglioli, E. H. (1883) Intorno a due nuovi pesci dal Golfo di Napoli. Zoologischer Anzeiger, 6, 397 - 400.","Nielsen, J. G. & Cohen, D. M. (1968) Redescription of Bellottia apoda Giglioli, 1883 (Pisces, Ophidioidea). Proceedings of the Linnean Society London, 179, 99 - 106.","Maul, G. E. (1976) The fishes taken in bottom trawls by R. V. \" Meteor \" during the 1967 Seamounts Cruise in the Northeast Atlantic. \" Meteor \" Forschungs - Ergebnisse, Reihe D, No. 22, 1 - 69.","Orsi, L. R. (1976) Fishes of Brotulidae family of Ligurian Sea. Rapport Commission internationale pour la Mer Mediterraee, 23 (8), 37 - 38.","Papaconstantinou, C. (1984) Occurrence of Bellottia apoda (Fam. Brotulidae) in the Greek seas. Cybium, 1984, 8 (2), 103 - 104.","Berdar, A., Giuffre, G., Cavallarro, G. & Potoschi, A. (1977) Contributo alla conoscenza del pesci spiaggiati lungo il littorale siciliano dello stretto di Messina. Memorie di Biologia Marina e di Oceanografia, 7 (5 - 6), 77 - 87.","Papaconstantinou, C., Tsimenidis, N. & Daulas, C. (1977) A new record of a bathypelagic fish Bellottia apoda Gigl., 1883 in Saronikos Gulf (Greece) (Pisces, Perciformes). Thalassographica, 1 (3), 279 - 287.","Jardas, I. (1979) Bellottia apoda Giglioli, 1883. A new genus and species in the Adriatic Sea. (Pisces, Perciformes, Brotulidae). Biljeske Institut za Oceanografiju i Ribarstvo, 38, 1 - 6.","Sardou, J, 1980: Contribution a la connaissance de la Faune Ichthyologique Liguro - Provencale: Bellottia apoda Giglioli, 1883 (Ophidioidei, Bythitidae), Poisson nouveau pour la Faune Francaise. Cybium 3 e ser., 1980 (9), 69 - 79.","Schwarzhans, W., 1981: Vergleichende morphologische Untersuchungen an rezenten und fossilen Otolithen der Ordnung Ophidiiformes. Berliner Geowissenschaftlische Abhandlungen, Reihe A, 32, 63 - 122.","de Ranieri, S. &. Sbrana, M. (1992) Presence de Bellottia apoda (Bythitidae) et de Epigonus constanciae (Apogonidae) dans la Mer Tyrrhenienne Septentrionale. Cybium 1992, 16 (2), 177 - 180.","Gramito, M. E. & Coen, B. (1997) New records of Bellottia apoda (Bythitidae) in the Adriatic Sea with notes on morphology and biology. Cybium 1997, 21 (2), 163 - 172.","Anon. (2002) Fish collection database of the American Museum of Natural History. American Museum of Natural History, Central Park West, NY 10024 - 5192, USA.","Mytilineou, C., Politou, C. - Y., Papaconstantinou C., Kavadas, S., D'Onghia, G. & Sion, L. (2005) Deep - water fish fauna in the Eastern Ionian Sea. Belgian Journal of Zoology, 135 (2), 229 - 233.","Nielsen, J. G. & Moller, P. R. (2008) New and rare deep - sea ophidiiform fishes from the Solomon Sea caught by the Danish Galathea 3 Expedition. Steenstrupia 30 (1), 21 - 46."]}

Nielsen, Jørgen G., Ross, Steve W., Cohen, Daniel M. (2009): Atlantic occurrence of the genus Bellottia (Teleostei, Bythitidae) with two new species from the Western North Atlantic. Zootaxa 2018: 45-57, DOI: 10.5281/zenodo.186025

Key to species of Bellottia spp. 1 a Long rakers on anterior gill arch 5���6............................................................................................................................ 2 1 b Long rakers on anterior gill arch 3���4............................................................................................................................ 4 2 a Caudal fin rays 6, precaudal vertebrae 10, total vertebrae 45, anal fin rays 74 ............ B. armiger (Philippine Islands) 2 b Caudal fin rays 7, precaudal vertebrae 10���12, total vertebrae 48���51, anal fin rays 75���88.......................................... 3 3 a Precaudal vertebrae 12, preopercular crest with spine (Fig. 3), depth at origin of anal fin 17.5 ���23.0 % of SL........... ........................................................................................................................................... B. robusta (Gulf of Mexico) 3 b Precaudal vertebrae 10, preopercular crest without spine, depth at origin of anal fin 17.0 % SL................................. .................................................................................................................................. B. cryptica (off southern Georgia) 4 a Dorsal fin rays 70���72, pectoral fin rays 19���20, origin of anal fin below dorsal fin ray no. 12���13, predorsal 44.0��� 45.5 % SL, spine on cleithrum present, no fangs on dentigerous bones................................................................................ ...................................................................................................... B. galatheae (Solomon Sea and Philippine Islands) 4 b Dorsal fin rays 83���94, pectoral fin rays 21���26, origin of anal fin below dorsal fin ray no. 17���19, predorsal 35.5 ���40.0 % SL, spine on cleithrum absent, all dentigerous bones with retrorse fangs................................................................. .................................................................................................................... B. apoda (Mediterranean and NE Atlantic), Published as part of Nielsen, J��rgen G., Ross, Steve W. & Cohen, Daniel M., 2009, Atlantic occurrence of the genus Bellottia (Teleostei, Bythitidae) with two new species from the Western North Atlantic, pp. 45-57 in Zootaxa 2018 on page 46, DOI: 10.5281/zenodo.186025

Bellottia robusta, new species (Figs. 1���4) Bellottia apoda: McEachran & Fechhelm (2005: 18) Bellotia sp.: Cordes et al. (2008: 783) Material examined (12 specimens, SL 43���82): Holotype: USNM 394117, SL 62, male, Gulf of Mexico, Green Canyon 234, 27�� 45 ���N, 91 �� 13 ���W, st. JSL- 4712, 535 m, coll. C. Fisher, 10 July 2004. Paratypes: TCWC 10956.01, SL 64, female, Gulf of Mexico, 27 �� 46.96 ���N, 91 �� 30.46 ���W, st: BH AT 1, 98 GOM JSL 4035, 540 ��� 580 m, coll. Tracy Ward, 7 Aug. 1998. - TCWC 10957.01, SL 53, female, Gulf of Mexico, 27 �� 44.7 ���N, 91 �� 13.3 ���W, st: GC AT 1, 98 GOM JSL 4033, 540 m, coll. Tracy Ward, 7 Aug. 1998. ��� USNM 394118, SL 34, male,Gulf of Mexico, Green Canyon 232, 27�� 44.49 ���N, 91 �� 19.07 ���W, st. JSL- 4437, 569 m, coll. C. Fisher, 24 June 2002. ��� USNM 394119, SL 35, male, Gulf of Mexico, Green Canyon 234, 27�� 44.78 ���N, 91 �� 13.30 ���W, st. JSL- 4569, 538 m, coll. C. Fisher, 26 Aug. 2003. ��� USNM 394120, SL 49, male, Gulf of Mexico, Garden Banks 543, 27�� 27.32 ���N, 93 �� 11.29 ���W, st. JSL- 4582, 546 m, coll. S. Hourdez, 3 Sep. 2003. ��� USNM 394121, SL 82, female, and ZMUC P 771652, female, Gulf of Mexico, same data as for holotype. ��� USNM 394122, SL 77, female, and ZMUC P 771653, SL 72, male, Gulf of Mexico, Green Canyon 234, 27�� 45.80 'N, 91 �� 13.30 ���W, st. JSL- 4713, 532 m, coll. C. Fisher, 10 July 2004. ��� USNM 394123, SL 50, male, Gulf of Mexico, Green Canyon 234, 27�� 44.76 ���N, 91 �� 13.46 ���W, st. JSL- 4720, 506 m, coll. K. Zelnio, 13 July 2004. ��� USNM 394124, SL 51, male, Gulf of Mexico, Green Canyon 234, 27�� 44.76 ���N, 91 �� 13.46 ���W, st. JSL- 4721, 506 m, coll. C. Fisher, 14 July 2004. Condition of material. In most specimens the body is much curled up and the mouth wide open making precise measurements difficult to obtain. Diagnosis. Bellottia robusta differs from its congeners by the following combination of characters: depth at origin of anal fin 17.5 ���23.0 % SL, small teeth blunt or pointed, four spines on hind margin and one on crest of preopercle, distinct spine on cleithrum above base of pectoral fin, anterior gill arch with 5���6 long rakers, predorsal 44.0��� 49.5 % SL and precaudal vertebrae 12. Similarity. Bellottia robusta is most similar to B. cryptica, in e. g. presence of a spine on cleithrum and in fin ray and total vertebral counts, but differs by the heavier body (17.5 ���23.0 vs 17.0 % SL at origin of anal fin), origin of anal fin below dorsal fin ray 10���15 vs 8, precaudal vertebrae 12 vs 10 and spine on preopercular crest vs no spine in B. cryptica. It differs from B. apoda by the deeper (17.0���23.0 vs 13.0���16.0 % SL) and darker body, more caudal fin rays (7 vs 6), poorly developed fangs (vs distinct, retrorse fangs), more long rakers on anterior gill arch (5���6 vs 3���4) and longer predorsal (44.0��� 49.5 vs 35.5 ���40.0 % SL). From B. galatheae it differs e. g. by the higher number of rays in dorsal (85 ��� 83 vs 70���72), caudal (7 vs 6), anal (75���88 vs 61) and pectoral (23���25 vs 19���20) fins. From B. armiger it differs by having more precaudal vertebrae (12 vs 10), more caudal fin rays (7 vs 6) and a larger predorsal length (44.0��� 49.5 vs 41.5 % SL). Description. The principal meristic and morphometric characters are shown in Table 1. The description is based mainly on the holotype. Differences of paratypes are given in brackets and in Table 1. B. robusta B. cryptica B. apoda Gulf of Mexico West. Atl. East. Atl. + Med. HT HT+ 11 paratypes Holotype n= 16 including LT Body robust, highest near base of pectoral fin, completely covered with oval, ca. 1.5 mm, cycloid scales. Anterior segment of lateral line extending close to dorsal margin of body and from above anus along midline of body. Head profile convex, cheek and gill-cover scaled, while rest of head naked. Mouth slightly oblique with posterior margin of maxillary vertically expanded ending well behind eye. Anterior nostril closer to upper lip than to posterior nostril, both with low rim. Small opercular spine strong and pointed, almost completely covered by skin, and not reaching posterior margin of opercle (a few specimens with spine reaching just beyond hind margin of opercle). Ventral margin of preopercle with four distinct spines and one spine on preopercular crest (Fig. 3) (spine often covered by skin; one specimen with two spines on crest). Origin of dorsal fin above tip of pectoral fin, anal fin origin just behind midpoint of fish (some specimens with preanal up to 59 % SL). Pectoral fin placed on mid-body level with peduncle higher than long. A distinct spine on cleithrum just above base of pectoral fin (one specimen with spine ending in a blunt tip). Anterior gill arch with 2���3 (2���4) plate-like rakers on upper branch, one long raker in the angle between the two branches and lower branch with 4���5 long rakers, followed by seven (6���9) plate-like rakers. Longest gill filament slightly shorter than longest raker. Two minute pseudobranchial filaments. Head pores: Because of thin, often torn skin, pores are difficult to observe. A few, large pores, with three (2���3) in a vertical row behind eye, three below eye and four pores on mandible. Small whitish papillae spread over head. Dentition: Most teeth are small and blunt. Head of vomer slightly curved with one tooth row of ca. ten small, blunt teeth. Posterior half of palatines edentate, and anterior half with one row of small, blunt teeth. Dentaries with posterior fifth edentate and dentigerous part with one tooth row increasing to 2���3 rows at symphysis; outer row with retrorse fangs. Premaxillaries with posterior third edentate and dentigerous part with one tooth row increasing to 4���5 rows at symphysis; outer row with larger, pointed teeth (three specimens with larger, pointed teeth on all dentigerous bones). Axial skeleton (based on radiographs): Tips of neural and haemal spines thin and pointed. Anterior neural spine one third the length of second spine. Neural spines 3���7 (2���7) slightly depressed. Bases of neural spines 4���7 (4���8) enlarged. Parapophyses present on vertebrae 8���12 (7���12) and pleural ribs on 4���10 (3���11). Epipleural ribs indistinct. Otolith (Fig. 2): The 3.5 mm elongate sagittal otolith (from a 64 mm SL paratype, TCWC 10956.01) is twice as long as high and twice as high as thick. Dorsal and ventral rim roundish, vaguely pointed posteriorly. Sulcus undivided, 1 / 3 ��� 1 / 4 the length of the otolith. No osteal channel. Coloration: Fig. 1 shows the holotype after four years of preservation. The body is brownish with light vertical fins. The abdomen, operculum and eye-surroundings dark blue and the rest of the head colored like the body. The body of some of the smaller paratypes is dark brown. Biology. This species appears to be very closely associated with complex habitat, provided by tubeworms or deep-sea corals (e.g., Lophelia pertusa). Rocky substrata are also likely habitats for B. robusta, even though it has not yet been collected there. This species is well concealed within crevices provided by these habitats, and specimens were not observed prior to collection. Because of its cryptic nature there are no more data on the biology or ecology of this species. A 53 mm specimen (TCWC 10957.01) contained eggs up to 0.7 mm in diameter; embryos were not observed. Etymology. The specific name, robusta, refers to the short, deep body. Distribution (Fig. 4). Known from nine localities in a restricted area of the northern Gulf of Mexico all caught by JSL at depths of 506��� 580 m., Published as part of Nielsen, J��rgen G., Ross, Steve W. & Cohen, Daniel M., 2009, Atlantic occurrence of the genus Bellottia (Teleostei, Bythitidae) with two new species from the Western North Atlantic, pp. 45-57 in Zootaxa 2018 on pages 47-51, DOI: 10.5281/zenodo.186025, {"references":["McEachran, J. D. & Fechhelm, J. D. (2005) Fishes of the Gulf of Mexico. Volume 2: Scorpaeniformes to Tetraodontiformes. University of Texas Press, Austin, 1004 pp.","Cordes, E. E., McGinley, M. P., Podowski, E. L., Becker, E. L., Lessard - Pilon, S., Viada, S. T. & Fisher, C. R. (2008) Coral communities of the deep Gulf of Mexico. Deep - Sea Research I. 55, 777 - 787."]}

Bellottia robusta, new species (Figs. 1–4) Bellottia apoda: McEachran & Fechhelm (2005: 18) Bellotia sp.: Cordes et al. (2008: 783) Material examined (12 specimens, SL 43–82): Holotype: USNM 394117, SL 62, male, Gulf of Mexico, Green Canyon 234, 27° 45 ’N, 91 ° 13 ’W, st. JSL- 4712, 535 m, coll. C. Fisher, 10 July 2004. Paratypes: TCWC 10956.01, SL 64, female, Gulf of Mexico, 27 ° 46.96 ’N, 91 ° 30.46 ’W, st: BH AT 1, 98 GOM JSL 4035, 540 – 580 m, coll. Tracy Ward, 7 Aug. 1998. - TCWC 10957.01, SL 53, female, Gulf of Mexico, 27 ° 44.7 ’N, 91 ° 13.3 ’W, st: GC AT 1, 98 GOM JSL 4033, 540 m, coll. Tracy Ward, 7 Aug. 1998. – USNM 394118, SL 34, male,Gulf of Mexico, Green Canyon 232, 27° 44.49 ’N, 91 ° 19.07 ’W, st. JSL- 4437, 569 m, coll. C. Fisher, 24 June 2002. – USNM 394119, SL 35, male, Gulf of Mexico, Green Canyon 234, 27° 44.78 ’N, 91 ° 13.30 ’W, st. JSL- 4569, 538 m, coll. C. Fisher, 26 Aug. 2003. – USNM 394120, SL 49, male, Gulf of Mexico, Garden Banks 543, 27° 27.32 ’N, 93 ° 11.29 ’W, st. JSL- 4582, 546 m, coll. S. Hourdez, 3 Sep. 2003. – USNM 394121, SL 82, female, and ZMUC P 771652, female, Gulf of Mexico, same data as for holotype. – USNM 394122, SL 77, female, and ZMUC P 771653, SL 72, male, Gulf of Mexico, Green Canyon 234, 27° 45.80 'N, 91 ° 13.30 ’W, st. JSL- 4713, 532 m, coll. C. Fisher, 10 July 2004. – USNM 394123, SL 50, male, Gulf of Mexico, Green Canyon 234, 27° 44.76 ’N, 91 ° 13.46 ’W, st. JSL- 4720, 506 m, coll. K. Zelnio, 13 July 2004. – USNM 394124, SL 51, male, Gulf of Mexico, Green Canyon 234, 27° 44.76 ’N, 91 ° 13.46 ’W, st. JSL- 4721, 506 m, coll. C. Fisher, 14 July 2004. Condition of material. In most specimens the body is much curled up and the mouth wide open making precise measurements difficult to obtain. Diagnosis. Bellottia robusta differs from its congeners by the following combination of characters: depth at origin of anal fin 17.5 –23.0 % SL, small teeth blunt or pointed, four spines on hind margin and one on crest of preopercle, distinct spine on cleithrum above base of pectoral fin, anterior gill arch with 5–6 long rakers, predorsal 44.0– 49.5 % SL and precaudal vertebrae 12. Similarity. Bellottia robusta is most similar to B. cryptica, in e. g. presence of a spine on cleithrum and in fin ray and total vertebral counts, but differs by the heavier body (17.5 –23.0 vs 17.0 % SL at origin of anal fin), origin of anal fin below dorsal fin ray 10–15 vs 8, precaudal vertebrae 12 vs 10 and spine on preopercular crest vs no spine in B. cryptica. It differs from B. apoda by the deeper (17.0–23.0 vs 13.0–16.0 % SL) and darker body, more caudal fin rays (7 vs 6), poorly developed fangs (vs distinct, retrorse fangs), more long rakers on anterior gill arch (5–6 vs 3–4) and longer predorsal (44.0– 49.5 vs 35.5 –40.0 % SL). From B. galatheae it differs e. g. by the higher number of rays in dorsal (85 – 83 vs 70–72), caudal (7 vs 6), anal (75–88 vs 61) and pectoral (23–25 vs 19–20) fins. From B. armiger it differs by having more precaudal vertebrae (12 vs 10), more caudal fin rays (7 vs 6) and a larger predorsal length (44.0– 49.5 vs 41.5 % SL). Description. The principal meristic and morphometric characters are shown in Table 1. The description is based mainly on the holotype. Differences of paratypes are given in brackets and in Table 1. B. robusta B. cryptica B. apoda Gulf of Mexico West. Atl. East. Atl. + Med. HT HT+ 11 paratypes Holotype n= 16 including LT Body robust, highest near base of pectoral fin, completely covered with oval, ca. 1.5 mm, cycloid scales. Anterior segment of lateral line extending close to dorsal margin of body and from above anus along midline of body. Head profile convex, cheek and gill-cover scaled, while rest of head naked. Mouth slightly oblique with posterior margin of maxillary vertically expanded ending well behind eye. Anterior nostril closer to upper lip than to posterior nostril, both with low rim. Small opercular spine strong and pointed, almost completely covered by skin, and not reaching posterior margin of opercle (a few specimens with spine reaching just beyond hind margin of opercle). Ventral margin of preopercle with four distinct spines and one spine on preopercular crest (Fig. 3) (spine often covered by skin; one specimen with two spines on crest). Origin of dorsal fin above tip of pectoral fin, anal fin origin just behind midpoint of fish (some specimens with preanal up to 59 % SL). Pectoral fin placed on mid-body level with peduncle higher than long. A distinct spine on cleithrum just above base of pectoral fin (one specimen with spine ending in a blunt tip). Anterior gill arch with 2–3 (2–4) plate-like rakers on upper branch, one long raker in the angle between the two branches and lower branch with 4–5 long rakers, followed by seven (6–9) plate-like rakers. Longest gill filament slightly shorter than longest raker. Two minute pseudobranchial filaments. Head pores: Because of thin, often torn skin, pores are difficult to observe. A few, large pores, with three (2–3) in a vertical row behind eye, three below eye and four pores on mandible. Small whitish papillae spread over head. Dentition: Most teeth are small and blunt. Head of vomer slightly curved with one tooth row of ca. ten small, blunt teeth. Posterior half of palatines edentate, and anterior half with one row of small, blunt teeth. Dentaries with posterior fifth edentate and dentigerous part with one tooth row increasing to 2–3 rows at symphysis; outer row with retrorse fangs. Premaxillaries with posterior third edentate and dentigerous part with one tooth row increasing to 4–5 rows at symphysis; outer row with larger, pointed teeth (three specimens with larger, pointed teeth on all dentigerous bones). Axial skeleton (based on radiographs): Tips of neural and haemal spines thin and pointed. Anterior neural spine one third the length of second spine. Neural spines 3–7 (2–7) slightly depressed. Bases of neural spines 4–7 (4–8) enlarged. Parapophyses present on vertebrae 8–12 (7–12) and pleural ribs on 4–10 (3–11). Epipleural ribs indistinct. Otolith (Fig. 2): The 3.5 mm elongate sagittal otolith (from a 64 mm SL paratype, TCWC 10956.01) is twice as long as high and twice as high as thick. Dorsal and ventral rim roundish, vaguely pointed posteriorly. Sulcus undivided, 1 / 3 – 1 / 4 the length of the otolith. No osteal channel. Coloration: Fig. 1 shows the holotype after four years of preservation. The body is brownish with light vertical fins. The abdomen, operculum and eye-surroundings dark blue and the rest of the head colored like the body. The body of some of the smaller paratypes is dark brown. Biology. This species appears to be very closely associated with complex habitat, provided by tubeworms or deep-sea corals (e.g., Lophelia pertusa). Rocky substrata are also likely habitats for B. robusta, even though it has not yet been collected there. This species is well concealed within crevices provided by these habitats, and specimens were not observed prior to collection. Because of its cryptic nature there are no more data on the biology or ecology of this species. A 53 mm specimen (TCWC 10957.01) contained eggs up to 0.7 mm in diameter; embryos were not observed. Etymology. The specific name, robusta, refers to the short, deep body. Distribution (Fig. 4). Known from nine localities in a restricted area of the northern Gulf of Mexico all caught by JSL at depths of 506– 580 m.

Bellottia Giglioli, 1883 Bellottia Giglioli, 1883: 399, type species Bellottia apoda Giglioli, 1883 by monotypy (type locality: Gulf of Naples, Italy). Xenobythites Smith & Radcliffe in Radcliffe, 1913, type species Xenobythites armiger Smith & Radcliffe in Radcliffe, 1913: 173 by original designation (type locality: Mindanao, Philippine Isls.) Diagnosis. The diagnosis in Nielsen et al. (1999: 97) is here slightly modified based on additional material. Body short, depth at origin of anal fin 13.0-23.0 % SL, head partly scaled, 3-5 spines at lower half of hind margin of preopercle, a pointed spine near upper angle of opercle, palatine teeth present, a few, large sensory pores behind and below eye and on mandible, long rakers on anterior gill arch 3-6, pectoral fin rays 19-28, pelvic fins absent, precaudal vertebrae 10-13, males with large genital hood. Similarity. The subfamily Bythitinae is characterized by continuous dorsal, caudal and anal fins and the penis being an integrated part of the large genital hood. In addition to Bellottia, two genera contain species that lack pelvic fins: Pseudonus Garman, 1899 and Hephthocara Alcock, 1892. The former of these is most similar to Bellottia with one species without and 1-2 species with pelvic fins, large sensory pores on head, a partly naked head and a very similar otolith. Hephthocara holds 1-2 species both without pelvic fins, but with no large sensory pores, a naked head and an otolith less similar to that of Bellottia. Species. Besides the five species mentioned in the introduction, Nielsen et al. (1999: 97) noted that two specimens collected in midwater (one from off Portugal and one from the Galapagos Islands) may represent undescribed species. Unfortunately, neither of the two specimens can be located., Published as part of Nielsen, J��rgen G., Ross, Steve W. & Cohen, Daniel M., 2009, Atlantic occurrence of the genus Bellottia (Teleostei, Bythitidae) with two new species from the Western North Atlantic, pp. 45-57 in Zootaxa 2018 on page 46, DOI: 10.5281/zenodo.186025, {"references":["Giglioli, E. H. (1883) Intorno a due nuovi pesci dal Golfo di Napoli. Zoologischer Anzeiger, 6, 397 - 400.","Radcliffe, L. (1913) Descriptions of seven new genera and thirty - one new species of fishes of the families Brotulidae and Carapidae from the Philippine Islands and the Dutch East Indies. Proceedings of the United States National Museum, 44, 135 - 176. Ross, S. W. & Quattrini, A. M. (2007) The fish fauna associated with deep coral banks off the southeastern United States. Deep-Sea Research, I, 54 (6), 975 - 1007.","Nielsen, J. G, Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) FAO species catalogue. Vol. 18. Ophidiiform fishes of the world (Order Ophidiiformes). An annotated and illustrated catalogue of pearlfishes, cusk - eels, brotulas and other ophidiiform fishes known to date. FAO Fisheries Synopsis. No. 125, vol. 18. Rome, FAO. 1999. 178 pp.","Garman, S. (1899) The fishes. Memoirs of the Museum of Comparative Zoology, 24, 1 - 431.","Alcock, A. (1892) Natural history notes from H. M. Indian Marine Survey Steamer \" Investigator \". - Ser. II, No. 5. On the bathybial fishes collected during the season of 1892 - 92. Annals and Magazine of Natural History, 6 (10), 345 - 365."]}